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The Middle Pleistocene archaeo-palaeontological site of Fontana Ranuccio (Anagni, central Italy) yielded a rich fau-nal assemblage consisting of more than 20,000 fossil remains, including four human teeth and multiple bone and lithic tools. The Ursus specimens from Fontana Ranuccio were historically ascribed to Ursus deningeri and Ursus arctos although a formal study has never been carried out. In this work, we have described for the first time all the Ursus material from Fontana Ranuccio reassessing the taxonomical attributions and discuss the first occurrence of the brown bear in the Italian Peninsula. Biometrical and morphological analyses confirm the presence of Ursus deningeri but, due the scarcity of diagnostic features and/or their preservation , some isolated specimens have been ascribed to Ursus sp. The results of this work allow to reconsider the earliest occurrence of U. arctos, attested in the Italian Peninsula to the late Middle Pleistocene site of Bucine.
Jacopo Conti1,2, Luca Bellucci3,4, Dawid A. Iurino2,
Flavia Strani2,4, Raffaele Sardella2,4
1 Polo museale, Sapienza Università di Roma, Italy.
2 Dipartimento di Scienze della Terra, PaleoFactory, Sapienza Università di Roma, Italy.
3 Museo di Geologia e Paleontologia, Sistema Museale di Ateneo, Università degli Studi di Firenze, Italy.
4 Istituto Italiano di Paleontologia Umana, Roma, Italy.
Corresponding author: J. Conti <>
ABSTRACT: The Middle Pleistocene archaeo-palaeontological site of Fontana Ranuccio (Anagni, central Italy) yielded a rich fau-
nal assemblage consisting of more than 20,000 fossil remains, including four human teeth and multiple bone and lithic tools. The
Ursus specimens from Fontana Ranuccio were historically ascribed to Ursus deningeri and Ursus arctos although a formal study
has never been carried out. In this work, we have described for the first time all the Ursus material from Fontana Ranuccio reas-
sessing the taxonomical attributions and discuss the first occurrence of the brown bear in the Italian Peninsula. Biometrical and
morphological analyses confirm the presence of Ursus deningeri but, due the scarcity of diagnostic features and/or their preserva-
tion, some isolated specimens have been ascribed to Ursus sp. The results of this work allow to reconsider the earliest occurrence
of U. arctos, attested in the Italian Peninsula to the late Middle Pleistocene site of Bucine.
Keywords: Ursus arctos, Ursus deningeri, Galerian, biochronology, systematics, evolution, carnivores.
Available online
ISSN (print): 2279-7327, ISSN (online): 2279-7335
Alpine and Mediterranean Quaternary, 34 (1), 2021, 55-68
In the Middle Pleistocene locality of Fontana Ra-
nuccio (0.4 Ma; Pereira et al., 2018), Ursus fossils are
represented by some isolated teeth and a third phalanx,
found during the 1980s excavation campaigns led by
researchers of the Istituto Italiano di Paleontologia Uma-
na (hereinafter IsIPU). These remains were firstly as-
signed to Ursus deningeri by Biddittu et al. (1979) and
Cassoli & Segre Naldini (1993) which described only
three specimens without providing a detailed compari-
son. Few years later, Azzaroli (1983) reported the pres-
ence of the brown bear Ursus arctos in the “Ranuccio
local fauna” without providing any description of the
fossil remains, thus opening the issue on the possible
co-occurrence of two bear species in the Fontana Ra-
nuccio site. Successively, Palombo et al. (2002 and
references therein) reported in their faunal list only the
presence of U. deningeri at Fontana Ranuccio, but with-
out performing any direct study of the fossil material. In
later publications, some authors reported the first occur-
rence (FO) of U. arctos at Fontana Ranuccio without
supporting this statement with any comparison or analy-
sis (Capasso Barbato et al., 1990; Di Stefano et al.,
1994; Sardella et al., 2006; Petronio et al., 2011). Final-
ly, in a recent publication, Petronio et al. (2019) consid-
ered the bear remains from Fontana Ranuccio as U.
deningeri, moreover indicating for this site the last occur-
rence (LO) of the species. As in the previous works, the
attribution was probably made on the basis of literature
data, since there is a lack of direct descriptions and
analyses of the materials. As a consequence of these
ambiguous attributions, the taxonomy of Ursus materials
from Fontana Ranuccio has been doubtful. Therefore,
on the basis of the above-mentioned literature as well as
from a biochronological point of view, the FO of the
brown bear or the LO of the Deninger bear in Italy, or
even both, could potentially be documented at Fontana
During the Pleistocene several fossil bear species
are recorded in the Italian Peninsula, such as the Au-
vergne bear (Ursus minimus, Devèze de Chabrol &
Bouillet, 1827), the Etruscan bear (Ursus etruscus, Cuvi-
er, 1823), the Deninger bear (Ursus deningeri, von
Reichenau, 1904), the cave bear (Ursus spelaeus,
Rosenmüller, 1794) and the brown bear (Ursus arctos,
Linnaeus, 1758) (Conti, 1954; Conti, 2019; Borselli et
al., 1980; Bon et al., 1991; Mazza & Rustioni, 1994;
Palombo et al., 2002; Petronio et al., 2007, 2020; Pe-
trucci & Sardella, 2009).
The first occurrence of U. deningeri in the Italian
Peninsula is recorded in the late Early Pleistocene sites
Conti J. et al.
Fig. 1 - Geographical location of the Middle Pleistocene site of Fontana Ranuccio (modified by Strani et al., 2018a).
of Slivia (Ambrosetti et
al., 1979; Palombo et
al., 2002 and reference
therein; Petronio et al.,
2011, 2019). Other
localities of the Early
and Middle Pleistocene
bearing U. deningeri
remains are Bristie
(Trentino Alto Adige;
Lugli & Sala, 2000),
Ponte Molle (Lazio;
Petronio et al., 2011;
Mecozzi et al., 2021),
Selva Vecchia (Veneto;
Bon et al., 1991), Iser-
nia La Pineta (Molise;
Peretto & Sala 2019),
Visogliano (Friuli Vene-
zia Giulia; Palombo et al., 2002 and reference therein)
and Venosa (Palombo et al., 2002 and reference the-
The oldest documented and figured specimen
attributed to Ursus arctos in the Italian Peninsula comes
from the late Middle Pleistocene site of Bucine (Upper
Valdarno, Tuscany) (Masini et al., 1991; Ferretti, 1997;
Gliozzi et al., 1997; Palombo et al., 2002). The material
consists in a fragmented cranium with the presence of
the upper fourth premolar and the first and second up-
per molars, now stored at the Museo di Geologia e
Paleontologia” at the Università di Firenze. The brown
bear is also reported in Sicily from two late Middle Pleis-
tocene/Late Pleistocene localities, Acquedolci and Con-
trada Camillà, both referred to the Elephas
mnaidriensis faunal complex” (Bonfiglio et al., 2001;
Pavia, 2001; Marra, 2003). The material is abundant but
fragmented, composed by three fragmentary hemi-
mandibles, isolated teeth and fragmentary postcranial
bones. Other fragmented bone elements (one indeter-
mined tooth and a phalanx) come from the layer 18
Ripardo del Poggio (Salerno), dated approximately to
MIS 6 (Boscato et al., 2009).
In this study, we present the first detailed descrip-
tion and comparative analysis of the bear specimens
from Fontana Ranuccio in order to: (1) define their taxo-
nomic status; (2) verify the possible co-occurrence of U.
arctos and U. deningeri at the site; (3) define the bio-
chronological aspects as the possible FO of U. arctos or
the LO of U. deningeri in the Italian Peninsula.
The archaeo-palaeontological site of Fontana Ra-
nuccio (Fig. 1) is located in the Anagni Basin (Biddittu et
al., 1979; Cassoli & Segre Naldini, 1993; Florindo et al.,
2021 and references therein), about 90 km southeast of
Rome, and it was discovered in 1976 during quarry ac-
tivities for the exploitation of “pozzolana” (volcanic ash).
Since then, researchers of the IsIPU have carried out
different field activities, in particular under the direction
of A. Segre (1978-2002), F. Parenti (2004-2018) and S.
Grimaldi (2019 - present).
The fossiliferous layer has been recently dated to
0.4 Ma (Pereira et al., 2018). The vertebrate collection
includes more than 20,000 specimens that are now
stored in the IsIPU depository (Anagni, Frosinone). The
large mammal assemblage have been attributed to 15
taxa (Biddittu et al., 1979; Cassoli & Segre Naldini,
1993; Strani et al., 2018a; Strani et al., 2018b; Strani et
al., 2019): Palaeoloxodon antiquus, Stephanorhinus sp.,
Equus cf. mosbachensis, Hippopotamus amphibius,
Dama clactoniana, Cervus elaphus eostephanoceros,
Praemegaceros sp., Bos primigenius, Sus scrofa ferus,
Panthera sp., Crocuta crocuta, Canis mosbachensis, cf.
Vulpes sp., Macaca sylvanus, Ursus arctos, Ursus
deningeri and Ursus sp. Small mammal remains are
scanty and have been recently ascribed by Bona & Stra-
ni (2021) to Microtus (Terricola) sp., Talpa sp., Elyomys
sp., Lepus sp., Erinaceus sp., Rodentia indet., cf. Glis
sp. Finally, Homo remains have been reviewed by Rubi-
ni et al. (2014) and are associated to several (more than
800) lithic and bone tools (Segre, 1984). Since the pio-
neering work of Azzaroli (1983) this faunal assemblage
is a reference for the Italian biochronological framework
(Gliozzi et al., 1997; Petronio et al., 2011; Petronio et al.,
2019 and references therein). Finally, Grimaldi et al.
(2020) carried out a techno-functional study of the entire
lithic assemblage identifying five groups of retouched
tools i.e., cutting tools, pointed tools, scrapers, notches
and denticulates.
Ursus material from Fontana Ranuccio consists of
12 isolated remains (Fig. 2) now stored at the IsIPU
laboratory in Anagni (Frosinone). This fossil material
was mainly found in the 1980s field activities, in particu-
lar in the 1982, 1984, 1985 and 1989 years. Morphologi-
cal analysis, descriptions and comparisons have been
provided taking into account the from different research-
ers (Ballesio, 1983; Torres, 1988; Capasso Barbato et
al., 1990; Argant, 1991; Mazza & Rustioni, 1994; Augu-
ste, 1995; Rabeder, 1999; Quiles, 2003; Meloro, 2007;
Wagner & Sabol, 2007; Wagner & Čermák, 2012). Mor-
phological data have been compared with specimens of
U. etruscus, U. deningeri, U. spelaeus and U. arctos
Review of Ursus material from Fontana Ranuccio (Middle Pleistocene, Central Italy)
Tab. 1 - List of selected specimens used for morphological comparison. Asterisks indicate the material
directly studied by the authors.
from selected European Pleistocene and Holocene lo-
calities (Tab. 1). The ontogenetic stage was inferred
according the scheme proposed by Stiner (1998).
The measurements of mesiodistal length and buc-
colingual width have been taken with digital caliper to
the nearest 0.1 mm on each fossil specimen (Tab. 2),
with the exception of the teeth FR 85-1 S1, FR84-1 and
the third phalanx (FR 84 inv 56576) due fragmentary
preservation. Dentognathic morphology and measure-
ments follow the terminology proposed by Rabeder
(1999) and Torres (1988).
Despite the high biometric variability of the teeth in
the Early and Middle Pleistocene bears, we performed
standard bivariate plots of width vs. length of M1, p4, m2
and m3 in order to compare our samples with speci-
mens from this time span in order to test possible bio-
metrical patterns among the Fontana Ranuccio speci-
mens and other Pleistocene ursid taxa (Torres, 1984;
Capasso Barbato et al., 1990; Marra, 2003; Rabeder et
al., 2010; Wagner et al., 2017 and personal measure-
ments; Fig. 3).
Class Mammalia Linnaeus, 1758
Order Carnivora Bowdich, 1821
Family Ursidae Fisher [De Waldheim], 1817
Genus Ursus Linnaeus, 1758
Ursus sp.
Referred material: I2 right (FR Sd-2), i3 right (FR
85-2 P3), i3 left (FR 82-2 Res), c right (FR 85-1 S1), p4
right (FR 89-1), m1 left (FR 96-33), m2 left (FR 84-1), ph
-3 (FR 84 Inv 56576).
Incisors and fragmented material: the incisors,
except FR Sd-2, are well preserved and show the typical
Ursus morphology with a developed central cusp and
the presence of two outlined lateral lobes; the distal one
is less developed than the mesial one (Fig. 2 f-g-h). The
lower canine (FR 85-1 S1), the second lower molar (FR
84-1) and the first lower molar (FR 96-33) are fragment-
ed; therefore, it is not possible to acquire any linear
measurement or identify any distinctive character to
define a specific attribution (Fig. 2 i-l-m). However, it is
possible to ascribe the specimen FR 96-33 to a prime
adult individual (class VII). The third phalanx (FR 84 inv
56576, Fig. 2 n) shows a rather elongated morphology,
although it is particularly worn. The proximal portion,
which is articulated with the trochlea of the second phal-
anx, is semicircular and the dorsal and plantar swelling
that characterize this anatomical portion of the bone
element are poorly developed. The ungual portion is thin
with a gentle dorsal curvature until the most distal por-
tion, where it sharply curves. Due to high wear and the
lack of any discriminant characters, we ascribe these
remains to Ursus sp.
Lower fourth premolar: the lower fourth premolar
(FR 89-1) has a very simple morphology, with narrow
and elongated external profile. The protoconid is well
developed and undivided, unlike the metaconid and the
paraconid which are small and poorly developed. The
presence of a tubercled area in the occlusal surface is
not observed except for one small accessory cusp on
Conti J. et al.
Tab. 2 - List and Measures of Ursus material from the Fontana Ranuccio deposit (Measures in mm).
Quiles, 2003; Madurell-Malapeira et al., 2009; Rabeder
et al., 2010; Wagner & Čermák, 2012; Arsuaga et al.,
2014; Prat-Vericat et al., 2020).
This simpler morphology is related to the early
Pleistocene Etruscan bear U. etruscus, characterized by
the lack of accessory cusps or tubercled structures in
the chewing surface of molariforms, and considered as
ancestor of both lineages (Wagner & Sabol, 2007). The
teeth of the Late Pleistocene cave bear (Ursus ex gr.
spelaeus) show instead a higher complexity of the chew-
ing surface (Fig. 4 k-l), contrary to what is observed in
the Late Pleistocene and extant brown bears, which
possess a simpler and secodont morphology of the oc-
clusal surface through time (Fig. 4 m-n-o-p). These dif-
ferences are probably related with the dietary habits of
the two bear lineages, one omnivorous (U. arctos) and
the other mostly herbivorous (U. spelaeus) (Terlato et
al., 2018). Biometric analyses did not allow any specific
discrimination regarding the Middle Pleistocene speci-
mens (Fig. 3 b), while the dimensional range of U.
the lingual portion of the protoconid slope. All these fea-
tures occur in the arctoid tooth morphology and can be
found also in the Late Pleistocene and recent Ursus
arctos (Reynolds, 1906; Torres, 1988; Capasso Barbato
et al., 1990; Mazza & Rustioni, 1994; Quiles, 2003; Me-
loro, 2007; Rabeder et al., 2010). U. deningeri can dis-
play a higher morphological complexity of the occlusal
surface characterized by many tubercles and by a much
more elliptic external profile (Fig. 4 e-f-g-i), even if many
remains such as the early Middle Pleistocene speci-
mens from the site of Hundsheim (Austria, HH 305),
C718 Cave (Czech Republic, Rv 20003) (Fig. 4 d-h),
and some other specimens recorded from Sima de los
Huesos (Spain, Middle Pleistocene, ca. 0.35 Ma),
Caune de l’Arago, (France, Middle Pleistocene, ca. 0.50
Ma), Cal Guardiola (Spain, latest Early Pleistocene, ca.
1.2-0.86 Ma), display a p4 without any accessory cus-
pids and simpler morphology, remarking the high varia-
bility and the possible presence of ancestral characters
displayed by U. deningeri (García et al., 1997, 2007;
Review of Ursus material from Fontana Ranuccio (Middle Pleistocene, Central Italy)
Fig. 2 - Ursus material from Fontana Ranuccio in occlusal and lateral views: a) M1 right (FR 06-411), b) FR M1 left (FR 56574), c) p4 right
(FR 89-1), d) m2 left (FR sd-1), e) m3 right (FR 56575), f) i3 left (FR 82-2 Res), g) i3 right (FR 85-2 P3), h) I2 right (FR Sd-2), i) c right (FR
85-1 S1), l) m2 left (FR 84-1), m) m1 left (FR 96-33), n) III phalanx right (FR 84 inv 56576).
Conti J. et al.
Fig. 3 - Comparison of the length and width scatter plots of various Early and Middle Pleistocene species of the genus Ursus. a) M1; b) p4;
c) m2; d) m3.
deningeri and U. arctos strongly overlap (also due to the
lack of sufficient data of the Early/Middle Pleistocene
Ursus arctos). As mentioned above, a simple morpholo-
gy, with the lack of accessory cusps in the lower p4 is a
shared character between Ursus arctos and Ursus
deningeri, specially during the late Early-Middle Pleisto-
cene (Wagner et al., 2017). Despite the presence of
Ursus deningeri in the deposit, the large variability of the
morphology and biometric data do not allow a specific
attribution of the isolated p4. To be more conservative
we ascribe FR 98-1 to Ursus sp.
Ursus deningeri von Reichenau, 1904
Referred material: M1 left (FR 56574), M1 right
(FR 06-411), m2 left (FR sd-1), m3 right (FR 56575).
Upper first molars: the specimen FR 56574 (Fig. 2
b) shows sub-quadrangular profile with a non-rectilinear
protocone-hypocone line and the presence of some
tubercled structures on the talon surface. The central
constriction of the tooth is rather marked especially in
the lingual portion and the main cusps are well devel-
oped. The biometric data show, as expected, an overlap
between the deningeroid and arctoid specimens; despite
this, we can observe that FR 56574 falls within the di-
mensional range mostly occupied by the Early-Middle
Pleistocene U. deningeri (Fig. 3 a) and showing a rather
large size of this tooth compared to the average dimen-
sion of U. arctos.
The specimen FR 06-411 (Fig. 2 a) has some dif-
ferent features from FR 56574. The metastyle and the
parastyle are well developed, the metastyle more than
the parastyle, and the talon shows a tubercled surface,
more than FR 56574. On the other hand, the tooth has a
quadrangular shape, the distal and proximal area show
the same profile and the protocone-hypocone line ap-
pears quite straight. Biometric analyses indicate that the
specimen is placed at the limit of the average dimen-
sional range of U. arctos, falling within the variability of
U. deningeri (Fig. 3 a). The occlusal surface of both
teeth displays tubercles, a common speloid feature
(Capasso Barbato et al., 1990; Quiles, 2003). According
to Torres (1984) and Capasso Barbato et al. (1990) the
straight protocone-hypocone line of FR 06-411 should
be related to the arctoid lineage. The latter assumption
stands up against the morphological comparison here
proposed (Fig. 5), in fact specimens of U. deningeri from
Hundsheim (Fig. 5 e-f) show the same straight proto-
ipoconid line, more than brown bear, which instead pre-
sents a consistent variability for this character. Consid-
ered all these morphological and biometric features, we
ascribe these specimens to U. deningeri.
Second lower molar: The specimen FR sd-1 (Fig. 2
d) is characterized by a high complexity of the occlusal
surface with the presence of
numerous tubercled struc-
tures and accessory cusps,
well representing the typical
cave bear morphology (Fig.
6) consisting of a metaconid
d iv i d e d i n t o t h r e e
metastylids, typical of U.
deningeri and U. ex gr. spe-
laeus (Capasso Barbato et
al., 1990; Quiles, 2003;
Torres, 1988). The biometric
analysis still indicates a
straight dimensional overlap
between U. deningeri and U.
arctos, which does not allow
to distinguish the two species
based on size. The specimen
FR SD-1 falls perfectly in this
overlap range (Fig. 3 c). Con-
cerning the morphological
features of the m2 from Fon-
tana Ranuccio we ascribe
th i s specimen t o U .
Lower third molar: The
specimen FR 56575 shows a
quadrangular outline with a
rounded talonid. The occlusal
surface is highly tubercled,
typical of the speloid lineage
(Capasso Barbato et al.,
1990; Quiles, 2003; Torres,
1988) (Fig. 7). The tooth
shows low worn structures
and a well-developed ento-
conid and metaconid, the
latter is composed by two
distinct cusps. Biometric data
also suggest the affinity of
the specimen with the
speloid group, fitting within
the dimensional range of U.
deningeri (Fig. 3 d). Accord-
ing to both morphological
and biometric analyses, we
ascribe this specimen to U.
The presence of U.
deningeri is confirmed at
Fontana Ranuccio, as al-
ready proposed by Biddittu et al., (1979) and Cassoli &
Segre Naldini, (1993). This species is widely distributed
throughout Europe and quite well represented in Italy
during the Middle Pleistocene (Fig. 8). U. deningeri was
a medium-large sized bear generally deemed as the
ancestor of Ursus spelaeus s.l. The transition between
these two species occurred during the Middle - Late
Pleistocene transition, so far these taxa are considered
two chronospecies, phylogenetically closely related
(Sardella et al., 2006, Valdiosera et al., 2007). From
molecular studies a complex scenario emerges during
the Late Pleistocene, with three different evolutionary
lines: U. spelaeus, U. ingressus and U. deningeri
kudarensis (Knapp et al., 2009). Late Pleistocene cave
bear in particular retained the same character of the
Middle Pleistocene Deninger bear, even if more marked,
Review of Ursus material from Fontana Ranuccio (Middle Pleistocene, Central Italy)
Fig. 4 - Comparison of the p4: a) Ursus arctos (left), DA-4B 18-36 (Deutsch-Altenburg, Early Pleisto-
cene); b) Ursus etruscus (right, mirrored in the table), IGF 911 (Valdarno, Early Pleistocene); c) Ursus
etruscus (left), IGF 4605 (Valdarno, Early Pleistocene); d) Ursus deningeri (left), HH 305 (Hundsheim,
early Middle Pleistocene); e) Ursus deningeri (left), HH 306 (Hundsheim, early Middle Pleistocene); f)
Ursus deningeri (left), Rv 20003 (Koněprusy Caves, early Middle Pleistocene); g) Ursus deningeri
(left), Rv 20005 (Koněprusy Caves, early Midde Pleistocene); h) Ursus deningeri (left), n.c. (Isernia la
Pineta, early Middle Pleistocene); i) Ursus deningeri (left), specimen H (Sandalja, Middle Pleistocene);
j) FR 89-1 (right, mirrored in the table) (Fontana Ranuccio, Middle Pleistocene); k) Ursus savini nor-
dostensis ssp. (left), IAM F-2365 (Ovrag, Late Pleistocene); l) Ursus spelaeus (left), P3021 (Caverna
delle Fate, Late Pleistocene); m) Ursus arctos (left), ZIN 34595 (Kudaro 3, Late Pleistocene); n) Ursus
arctos (left), V.1152 (Vigna S. Carlo, Late Pleistocene); o) Ursus arctos (left), 3399 (Alps, Recent); p)
Ursus arctos (right, mirrored in the table), 3362 (Alps, Recent). Abbreviations: Med Metaconid, Prd
Protoconid, Pad Paraconid.
as the broad, domed, steep forehead, that result in a
‘step’ in the midsagittal plane (Santos et al., 2017).
During the Middle Pleistocene, U. arctos has been
reported in some Italian localities: Bucine (Upper Val-
darno, Tuscany; Mazza, 1998), Riparo del Poggio
(Salerno; Boscato et al., 2009), Acquedolci and Contra-
da Camillà (Sicily; Bonfiglio et al., 2001; Marra, 2003;
Pavia, 2001) (Fig. 8). The brown bear record from Ri-
paro del Poggio is composed only by two elements (one
indeterminate tooth and a phalanx), the deposit consists
in a shelter which was part of a complex underground
karst system, which was partially dismantled by sea
erosion. The Ursus arctos material come from the lower
level dated to the cold phase of MIS 6 (Boscato et al.,
2009). The cranium from Bucine comes from one of
many small fossiliferous deposits as Poggio Amaro, Le
Capannelle, Pogi di Bucine, Capannole, Migliorini, Cava
di Bucine, Cava Le Vigne, Cava del Rinoceronte, mostly
situated along the Ambra creek, a left-hand tributary of
the Arno river, and excavated since the 1940. No infor-
mation is available on the exact locality where the crani-
um of U. arctos was recovered, but all the fossil remains
are dated around the late Middle Pleistocene (referable
to the Saalian cycle, from 340 ky to 120 ky ago; Mazza,
Conti J. et al.
Fig. 5 - Comparison of the M1. a) FR 06-411 (right). b) FR
56574 (left, mirrored in the table). c) U. deningeri (left, mirrored
in the table), HH/5/342 (Hundsheim, Middle Pleistocene) d) U.
deningeri (left, mirrored in the table), HH/5/398, (Hundsheim,
Middle Pleistocene). e) U. arctos (left, mirrored in the table),
DA4B/18 (Deutsch-Altenburg, Early Pleistocene). f) U. arctos
(right), DA4V/14 (Deutsch-Altenburg, Early Pleistocene). g) U.
arctos priscus (right), W S.1 (Winden, Late Pleistocene). h) U.
arctos (left, mirrored in the table), C. 4 (Banskà Bystrica, Re-
cent). i) U. arctos (right), IGF 10961 (Bucine, late Middle Pleis-
tocene). e-f-g-c) modified from Rabeder et al. (2010).
Fig. 7 - Comparison of the m3. a) FR 56575 (right). b) U. arctos
(left, mirrored in the table), C.4, (Banská Bystrica, recent). c) U.
deningeri (left, mirrored in the table), MF/1346/45 (Kozi Grzbiet,
Middle Pleistocene). d) U. deningeri (left, mirrored in the table),
Hund 382 (Hundsheim, Middle Pleistocene).
Fig. 6 - Comparison of the m2. a) FR 84-1 (left). b) FR sd-1 (left).
c) U. arctos (right, mirrored in the table), 3364 (Alps, recent). d)
U. deningeri (right, mirrored in the table), 1889/5/404
(Hundsheim, Middle Pleistocene). e) U. deningeri (left),
MF/1346/37 (Kozi Grzbied, Middle Pleistocene), modified from
Wagner et al. (2012).
The bear remains from Sicily
have been excavated in the last forty
years, and have been found associat-
ed with Cervus elaphus siciliae and
Elephas mnaidriensis, dated to the late
Middle Pleistocene/Late Pleistocene
(E. mnaidriensis Faunal Complex, from
early MIS 6 to MIS 4 stages; Bonfiglio
et al., 2001, 2003). The occurrence of
the brown bear is also documented in
some other sites of the north/eastern
Italian Peninsula (see Bon et al.,
1991), however this material needs
further chronological and systematic
Since the uncertain dating of
these deposits, it is possible to hypoth-
esize that the Bucine specimen could
be the F.O. of the brown bear in the
Italian peninsula. A recent paleoenvi-
ronmental reconstruction based on
dietary adaptations of ungulates from
Fontana Ranuccio suggests that a
mosaic of both closed and open land-
scapes characterized this locality
around 0.4 Ma with a relative abun-
dance of soft plant resources and
probably affected by a marked season-
ality (Strani et al., 2018a; Strani et al.,
2019; Strani, 2020; Strani et al., 2021).
A heterogeneous environment could
have favored the coexistence of two
different bear species in the area,
which may have lived in sympatric
conditions exploiting different trophic
resources as largely documented in
several Late Pleistocene localities of
the Italian Peninsula (Capasso Barbato
et al., 1990; Minieri et al., 1995; Pa-
lombo et al., 2002; Mazza et al., 2005).
Although palaeoecological con-
siderations allow the presence of the
two species, paleontological data do
not show with any reliable evidence of
the presence of Ursus arctos. The
fossils from Fontana Ranuccio repre-
sent the LO of Ursus deningeri in Italy.
New excavations are still ongoing car-
ried on by IsIPU and new findings will
better address many unsolved ques-
tions and shed new light on the distri-
bution and evolution of bears in Europe.
We are very grateful to the Italian Institute of Hu-
man Paleontology for granting access to the material
and for the continuous assistance during the analyses.
We would like to express also gratitude to Martin Sabol
(Comenius University, Bratislava), for advising and the
inspiring discussion that made possible this work. We
also thank the MuSe (Museo delle Scienze of Trento),
the Parco Nazionale d’Abruzzo, Lazio e Molise, the
Department of Geology and Paleontology of Comenious
University (Bratislava, Slovakia), the Slovak Museum of
Nature Protection and Speleology (Liptovský Mikuláš,
Slovakia), the National History Museum (Wien, Austria),
the Department of Paleontology of Wien University
(Wien, Austria), for granting access to fossil and extant
bear material used for morphological comparisons. We
thank Chiara Delpino of Soprintendenza Archeologia
Belle Arti e Paesaggio per le Province di Frosinone,
Latina e Rieti (Ministero della Cultura) and the Italian
Institute of Human Paleontology for granting access to
Review of Ursus material from Fontana Ranuccio (Middle Pleistocene, Central Italy)
Fig. 8 - Distribution of the fossil bears during the Middle Pleistocene of the Italian Peninsu-
la. 1) Venosa (van Heteren et al., 2019). 2) Bristie I (Lugli & Sala, 2000). 3) Cava Rinaldi
(Petronio et al., 2019). 4) Cretone (Petronio et al., 2011). 5) Fontana Ranuccio (this work).
6) Isernia la Pineta (Peretto & Sala, 2019). 7) Ponte Molle (Petronio et al., 2011). 8) Slivia
(Bon et al., 1991; Petronio et al., 2011). 9) Valdemino (Ghezzo et al., 2015). 10) Visoglia-
no (Tozzi et al., 2000). 11) Cava Nord (Soave) (Rustioni & Mazza, 1993). 12) Cengelle 1
(Soave) (Rustioni & Mazza, 1993). 13) Castello (Soave) (Rustioni & Mazza, 1993). 14)
Cerè (Ghezzo et al., 2013; Rossi & Santi, 2011). 15) Selva Vecchia (van Heteren et al.,
2019). 16) Sorbano di Romagnano (Bon et al., 1991). 17) Fornace di Cornedo (Rustioni &
Mazza, 1993). 18) Grotta di San Bernardino (Bon et al., 1991). 19) Acquedolci (Marra,
2003). 20) Contrada Camilla (Marra, 2003). 21) Bucine (Mazza, 1997; Petronio et al.,
2019). 22) Castel di Guido (Caloi et al., 1998). 23) Prati Fiscali (Petronio et al., 2011). 24)
Fara Sabina (Petronio et al., 2011). 25) Riparo del Poggio (Boscato et al., 2009).
Conti J. et al.
the palaeontological collections of Fontana Ranuccio.".
We'd like to thank the two reviewers for providing useful
comments that allowed us to improve our manuscript.
JC acknowledges Sapienza, University of Rome for the
funds provided by the call “Bando per il finanziamento di
progetti di ricerca congiunti per la mobilità all’estero di
studenti di dottorato del XXXI e XXXII ciclo n.
2682/2017”, that allowed the acquisition of part of the
data present in this work.
Ambrosetti P., Bartolomei G., De Giuli C., Ficcarelli G.,
Torre D. (1979) - La breccia ossifera di Slivia
(Aurisina-Sistiana) nel Carso di Trieste. Bollettino
della Società Paleontologica Italiana, 18(2), 207-
Argant A. (1991) - Carnivores quaternaires en Bourgo-
gne. Doc. Lab. Géol. Lyon, 115, pp. 301.
Arsuaga J.L., Martínez I., Arnold L.J., Aranburu A.,
Gracia-Téllez A., Sharp W.D., Quam R.M.,
Falguères C., Pantoja-Pérez A., Bischoff J.,
Poza-Rey E., Parés J.M., Carretero M., Demuro
M., Lorenzo C., Sala N., Martinón-Torres M.,
García N., Alcázar de Velasco A., Cuenca-Bescós
G., Gómez-Olivencia A., Moreno D., Pablos A.,
Shen C., Rodríguez L., Ortega A.I., García R.,
Bonmatí A., Bermúdez de Castro J.M., Carbonell
E. (2014) - Neandertal roots: Cranial and chrono-
logical evidence from Sima de los Huesos.
Science, 344(6190), 1358-1363.
Auguste P. (1995) - Cadres biostratigraphiques et pa-
léoécologiques du peuplement humain dans la
France septentrionale durant le pléistocène: ap-
ports de l’étude paléontologique des grands mam-
mifères du gisement de Biache-Saint-Vaast (Pas-
de-Calais). Doctoral Dissertation, Paris, Muséum
National d’histoire Naturelle, pp. 1924.
Azzaroli A. (1983) - Quaternary mammals and the “End-
Villafranchian” dispersal event - A turning point in
the history of Eurasia. Palaeogeography, Palaeo-
climatology, Palaeoecology, 44(1-2), 117-139.
Ballesio R. (1983) - Le gisement Pléistocène supérieur
de la grotte de Jaurens à Nespouls (Corrèze) : les
Carnivores. Nouvelles Archives DuMuséum d’His-
toire Naturelle de Lyon, 21, 9-43.
Baryshnikov G.F. (2010) - Late Pleistocene brown bear
(Ursus arctos) from the Caucasus. Russian Jour-
nal of Theriology, 9(1), 9-17.
Biddittu I., Cassoli P.F., Radicati di Brozolo F., Segre
A.G., Segre Naldini E., Villa I. (1979) - Anagni a K/
Ar dated Lower Middle Pleistocene site, Central
Italy: preliminary report. Quaternaria, 21, 53-71.
Bon M., Piccoli G., Sala B. (1991) - I giacimenti quater-
nari di vertebrati fossili nell’Italia Nord-Orientale.
Mem. Se. Geol. Padova, 43, 185-231.
Bona F., Strani F. (2021) - New data on the Middle
Pleistocene small mammal fauna from the Homo
bearing site of Fontana Ranuccio (Anagni Basin,
Central Italy). Alpine and Mediterranean Quater-
nary, 34 (1), 69-74.
Bonfiglio L., Mangano G., Marra A.C., Masini F. (2001) -
A new Late Pleistocene vertebrate faunal complex
from Sicily (S. Teodoro Cave, North-Eastern Sicily,
Italy). Bollettino della Società Paleontologica Italia-
na, 2, 149-158.
Bonfiglio L., Di Maggio C., Marra A., Masini F., Petruso
D. (2003) - Bio-chronology of Pleistocene vertebra-
te faunas of Sicily and correlation of vertebrate
bearing deposits with marine deposits. Il Quaterna-
rio, 16(1), 107-114.
Borselli V., De Giuli C., Ficcarelli G., Mazzini M. (1980) -
Casa Frata: una località fossilifera del Villafran-
chiano Superiore presso Terranuova Bracciolini
(Arezzo) nel Valdarno Superiore. Bolletino della
Società Paleontologica Italiana, 19(2), 245-258.
Boscato P., Boschian G., Caramia F., Gambassini P.
(2009) - Il Riparo del Poggio a Marina di Camerota
(Salerno): culture ambiente. Rivista di scienze
preistoriche, 59(2009), 5-40.
Caloi L., Palombo M.R., Zarlenga F. (1998) - Late-
Middle Pleistocene mammal faunas of Latium
(central Italy): stratigraphy and environment. Qua-
ternary International, 47, 77-86.
Capasso Barbato L., Minieri M.R., Petronio C., Vigna
Taglianti A. (1990) - Strutture dentarie di Ursus
arctos e di Ursus spelaeus della grotta di Monte
Cucco (Sigillo, Perugia, Italia). Bollettino della So-
cietà Paleontologica Italiana, 29, 335-356.
Cassoli P.F., Segre Naldini E. (1993) - Le faune Villa-
franchiane: Costa san Giacomo e Fontana Aceto-
sa. Dives Anagnia. L’Erma Di Bretscheider, Roma,
Conti J. (2019) - Evolution of cranio-dental features and
distribution of brown bear (Ursus arctos L., 1758)
in Europe. Ph.D. Thesis. Sapienza University of
Rome, pp. 177.
Conti S. (1954) - Morfologia comparata craniale ed en-
cefalica degli orsi pleistocenici della Liguria. Me-
morie Del Museo Civico Di Storia Naturale “G.
Doria” Genova, pp. 86.
Cuvier G. (1823) - Recherches sur les Ossemens Fossi-
liles de quadrupédes. Paris, 5(1).
Devèze de Chabrol J.S., Bouillet J.B. (1827) - Essai
géologique et minéralogique sur les environs d'Is-
soire, département du Puy-de-Dôme, et principale-
ment sur la Montagne de Boulad, avec la descrip-
tion et les figures lithographiées des ossemens
fossiles qui y ont été recueillis. Ursidae. Thibaud-
Landriot, Clermont-Ferrand, pp. 140.
Di Stefano G., Petronio C., Sardella R. (1994) - Il signifi-
cato biocronologico e paleoecologico di alcuni taxa
di mammiferi del Plio-Pleistocene dell’Italia centra-
le. Studi Geologici Camerti, volume speciale 1994
“Biostratigrafia dell’Italia centrale”. Università degli
Studi di Camerino, Camerino, 459-467.
Ferretti M. (1997) - Mammuthus cf. primigenius
(Proboscidea, Mammalia), a new faunal element
from the late Middle Pleistocene of Conca river
(Cattolica, Romagna, Italy). Bollettino della Società
Paleontologica Italiana, 36(3), 391-398.
Florindo F., Marra F., Angelucci D.E., Biddittu I., Bruni
L., Florindo F., Gaeta M., Guillou H., Jicha B.,
Macrì P., Morigi C., Nomade S., Parenti F.,
Pereira A., Grimaldi S. (2021) - Environmental
evolution, faunal and human occupation since 2
Review of Ursus material from Fontana Ranuccio (Middle Pleistocene, Central Italy)
Ma in the Anagni basin, central Italy. Scientific
reports, 11(1), 7056.
García N., Arsuaga J.L., Torres T.D. (1997) - The carni-
vore remains from the Sima de los Huesos Middle
Pleistocene site (Sierra de Atapuerca, Spain).
Journal of Human Evolution, 33(2-3), 155-174.
García N., Santos E., Arsuaga J.L., Carretero J.M.
(2007) - Endocranial morphology of the Ursus
deningeri von Reichenau 1904 from the Sima de
los Huesos (Sierra de Atapuerca) Middle Pleisto-
cene site. Journal of vertebrate paleontology, 27
(4), 1007-1017.
Ghezzo E., Berté D., Sala B. (2013) - The revalutation
of Galerian Canidae, Felidae and Mustelidae of
the Ceré Cave (Verona, Northeastern Italy). Qua-
ternary International, 339-340, 76-89.
Ghezzo E., Boscaini A., Madurell-Malapeira J., Rook L.
(2015) - Lynx remains from the Pleistocene of
Valdemino cave (Savona, Northwestern Italy), and
the oldest occurrence of Lynx spelaeus
(Carnivora, Felidae). Rendiconti Lincei, 26(2), 87-
Gliozzi E., Abbazzi L., Argenti P., Azzaroli A., Caloi L.,
Capasso Barbato L., Di Stefano G., Esu D.,
Ficcarelli G., Girotti O., Kotsakis T., Masini F.,
Mazza P., Mezzabotta C., Palombo M.R., Petronio
C., Rook L., Sala B., Sardella R., Zanalda E. Torre
D. (1997) - Biochronology of selected mammals,
molluscs and ostracods from the Middle Pliocene
to the Late Pleistocene in Italy. The state of the
art. Rivista Italiana di Paleontologia e Stratigrafia,
103(3), 369-388.
Grimaldi S., Santaniello F., Angelucci D.E., Bruni L.,
Parenti F. (2020) - A Techno-Functional Interpre-
tation of the Lithic Assemblage from Fontana Ra-
nuccio (Anagni, Central Italy): an Insight into a
MIS 11 Human Behaviour. Journal of Paleolithic
Archaeology, 3, 944966.
Knapp M., Rohland N., Weinstock J., Baryshnikov G.,
Sher A., Nagel D., Rabeder G., Pinhasi R.,
Schmidt H.A., Hofreiter M. (2009) - First DNA se-
quences from Asian cave bear fossils reveal deep
divergences and complex phylogeographic pat-
terns. Molecular ecology, 18(6), 1225-1238.
Linnaeus C. (1758) - Systema naturæ per regna tria
naturæ, secundum classes, ordines, genera, spe-
cies, cum characteribus, differentiis, synonymis,
locis. 1 (10th ed.). Laurentius Salvius, Stockholm.
pp. 824.
Lugli C., Sala B. (2000) - La teriofauna del Pleistocene
medio di Bristie I°(Carso Triestino). Atti del Museo
Civico di Storia naturale di Trieste, 48, 35-58.
Madurell-Malapeira J., Alba D.M., Moyà-Solà S. (2009) -
Carnivora from the late Early Pleistocene of Cal
Guardiola (Terrassa, Vallès-Penedès Basin, Cata-
lonia, Spain). Journal of Paleontology, 83(6), 969-
Marra A.C. (2003) - Ursus arctos from selected Pleisto-
cene site of Eastern Sicily. Bollettino della Società
Paleontologica Italiana, 42(1-2), 145-150.
Masini F., Sala B., Ambrosetti P., Azzaroli A., Ficcarelli
G., Kotsakis T., Rook L., Torre D. (1991) - Mam-
malian faunas of selected villafranchian and gale-
rian localities. In: INQUA SEQS, Subcommission
for European Quaternary Stratigraphy, Cromer
Symposium, Norwich., UK, 3-7 September 1990.
Mazza P. (1998) - Taphonomic analysis of late Middle
Pleistocene mammal remains from Bucine. Bollet-
tino della Società Paleontologica Italiana, 36, 381-
Mazza P., Rustioni M. (1994) - On the phylogeny of
Eurasian bears. Palaeontographica, 230, 1-38.
Mazza P. (1997) - Taphonomic analysis of late Middle
Pleistocene mammal remains from Bucine. Bollet-
tino della Società Paleontologica Italiana, 36, 381-
Mazza P., Rustioni M., Agostini S., Rossi A. (2005) - An
unexpected Late Pleistocene macaque remain
from Grotta degli Orsi Volanti (Rapino, Chieti, cen-
tral Italy). Geobios, 38(2), 211-217.
Mecozzi B., Iannucci A., Mancini M., Sardella R. (2021)
- Redefining Ponte Molle (Rome, central Italy): an
important locality for Middle Pleistocene mammal
assemblages of Europe. Alpine and Mediterranean
Quaternary, 34(1), 131-154
Doi: 10.26382/AMQ.2021.09
Meloro C. (2007) - La fauna quaternaria di Grotta Milano
(Petina, Salerno). Atti I Convegno Regionale di
Speleologia “Campania Speleologica” 1-3 giugno
2007 Oliveto Citra (SA), pp. 75-83.
Minieri M., Petronio C., Sardella R., Scarano M. (1995) -
Le faune a mammiferi del Pleistocene superiore
dell’Italia peninsulare. Quaderni Padusa, 1, 75-87.
Palombo M.R., Azanza B., Alberdi M.T. (2002) - Italian
mammal biochronology from Latest Miocene to
Middle Pleistocene: a multivariate approach. Geo-
logica Romana, 36, 335-368.
Pavia M. (2001) - The Middle Pleistocene fossil avifauna
from the Elephas mnaidriensis Faunal Complex”
of Sicily (Italy): preliminary results. In Congres-
so Internazionale: "La Terra Degli Elefanti", Roma
16-20 ottobre 2001, 497-501.
Peretto C., Sala B. (2019) - Isernia La Pineta (Isernia).
Annali dell’Università di Ferrara. Sezione: Museo-
logia scientifica e naturalistica, 15, 31-38.
Pereira A., Nomade S., Moncel M.H., Voinchet P.,
Bahain J.J., Biddittu I., Falguères C., Giaccio B.,
Manzi,G., Parenti F., Scardia G., Scao V., Sottili
G., Vietti A. (2018) - Integrated geochronology of
Acheulian sites from the southern Latium (central
Italy): Insights on human-environment interaction
and the technological innovations during the MIS
11-MIS 10 period. Quaternary Science Reviews,
187, 112-129.
Doi: 10.1016/j.quascirev.2018.03.021
Petronio C., Di Canzio E., Salari L. (2007) - The Late
Pleistocene and Holocene Mammals in Italy: new
biochronological and paleoenvironmental data.
Palaeontographica, Abt. A, 279, 147-157.
Petronio C., Bellucci,L., Martinetto E., Pandolfi L., Salari
L. (2011) - Biochronology and palaeoenvironmen-
tal changes from the Middle Pliocene to the Late
Pleistocene in Central Italy. Geodiversitas, 33(3),
Petronio C., Di Stefano G., Kotsakis T., Salari L., Marra
F., Jicha B.R. (2019) - Biochronological framework
Conti J. et al.
for the late Galerian and early-middle Aurelian
Mammal Ages of peninsular Italy. Geobios, 53, 35-
Petronio C., Angelone C., Atzori P., Famiani F.,
Kotsakis T., Salari L. (2020) - Review and new
data of the fossil remains from Monte Peglia (late
Early Pleistocene, Central Italy). Rivista Italiana di
Paleontologia e Stratigrafia, 126 (3), 791-819.
Petrucci M., Sardella R. (2009) - Ursus etruscus Cuvier,
1823 from the Early Pleistocene of Monte Argen-
tario (Southern Tuscany, Central Italy). Bollettino
della Società Paleontologica Italiana, 48(2), 89-94.
Prat-Vericat M., Rufí I., Llenas M., Madurell-Malapeira J.
(2020) - Middle Pleistocene Ursus deningeri from
Grotte de la Carrière (Réseau Lachambre, Têt
Valley, Eastern Pyrenees). Journal of Iberian Geo-
logy, 46(2), 163-175.
Quiles J. (2003) - Les Ursidae du Pléistocène moyen et
supérieur en Midi méditerranéen: apports paléon-
tologiques, biochronologiques et archéozoolo-
giques. Doctoral Dissertation, Paris, Muséum Na-
tional d’histoire Naturelle, pp. 1307.
Rabeder G. (1999) - Die Evolution des Höhlenbärenge-
bisses. Verlag Der Österr. Akad. Der Wiss., 11, 1-
Rabeder G., Pacher M., Withalm G. (2010) - Early Plei-
stocene bear remains from Deutsch-Altenburg
(Lower Austria). Mitteilungen Der Kommission Für
Quartärforschung Der Österreichischen Akademie
Der Wissenschaften, 17, 1-135.
von Reichenau W. (1904) - Beiträge zur näheren Kennt-
niss der Carnivoren aus den Sanden von Mauer
and Mosbach. Abhandlungen der Grossherzolich
Hessischen geologischen Landesanstalt zu Darm-
stadt. 4(2), 14.
Reynolds S.H. (1906) - Monograph of British Mammalia
of the Pleistocene period. The Bears. London Pal-
aeontographical Society, 2, 1-35.
Rosenmüller J. (1794) - Quaedam de Ossibus Fossili-
bus Animalis cuiusdam, Historiam eius et Cogni-
tionem accurationrem illustrantia. Thesis, Leipzig,
pp. 34.
Rossi M., Santi G. (2011). Ursus deningeri-spelaeus
group from Cerè Cave (Veneto, North Italy) in the
new evolutionary frame of the cave bear. Part one:
skulls and mandibles. Annalen des Naturhistor-
ischen Museums in Wien. Serie A für Mineralogie
und Petrographie, Geologie und Paläontologie,
Anthropologie und Prähistorie. Naturhistorischen
Museum, Wien, 567-590.
Rubini M., Cerroni V., Festa G., Sardella R., Zaio P.
(2014) - A revision of hominin fossil teeth from
Fontana Ranuccio (Middle Pleistocene, Anagni,
Frosinone, Italy). Journal of Human Evolution, 77,
Doi: 10.1016/j.jhevol.2014.09.002
Rustioni M., Mazza P. (1993) - The Tibetan-like bear
from Grotta di Reale, Porto Azzurro (Isle of Elba,
Italy). Il Quaternario, 6(1), 35-38.
Santos E., Gómez-Olivencia A., Arlegi M., Arsuaga J.L.
(2017) - Cranial morphological differences within
U. deningeri - U. spelaeus lineage: A double tradi-
tional and geometric morphometrics approach.
Quaternary International. 433, 347-362.
Sardella R., Palombo M.R., Petronio C., Bedetti C.,
Pavia M. (2006) - The early Middle Pleistocene
large mammal faunas of Italy: an overview. Qua-
ternary International, 149(1), 104-109.
Segre A.G. (1984) - Considerazioni sulla cronostratigra-
fia del Pleistocene laziale. Atti Della XXIV Riunione
Scientifica Dell’Istituto Italiano Di Preistoria e Pro-
tostoria, Istituto Italiano Di Preistoria e Protostoria,
Firenze, 149-154.
Sher A. V., Weinstock J., Baryshnikov G. F., Davydov S.
P., Boeskorov G. G., Zazhigin V. S., Nikolskiy P.
A. (2011). The first record of “spelaeoid” bears in
Arctic Siberia. Quaternary Science Reviews, 30(17
-18), 2238-2249.
Stiner M.C. (1998) - Mortality analysis of Pleistocene
bears and its paleoanthropological relevance.
Journal of Human Evolution, 34(3), 303-326.
Strani F. (2020) - Impact of Early and Middle Pleisto-
cene major climatic events on the palaeoecology
of Southern European ungulates. Historical Biolo-
gy, in press.
Doi: 10.1080/08912963.2020.1782898
Strani F., DeMiguel D., Bona F., Sardella R., Biddittu I.,
Bruni L., De Castro A., Guadagnoli F., Bellucci L.
(2018a) - Ungulate dietary adaptations and palae-
oecology of the Middle Pleistocene site of Fontana
Ranuccio (Anagni, Central Italy). Palaeogeo-
graphy, Palaeoclimatology, Palaeoecology, 496,
Doi: 10.1016/j.palaeo.2018.01.041
Strani F., Profico A., Manzi G., Pushkina D., Raia P.,
Sardella R., DeMiguel D. (2018b) - MicroWeaR: a
new R package for dental microwear analysis.
Ecology and Evolution, 8, 7022-7030.
Strani F., Pushkina D., Bocherens H., Bellucci L.,
Sardella R., DeMiguel D. (2019) - Dietary Adapta-
tions of Early and Middle Pleistocene Equids From
the Anagni Basin (Frosinone, Central Italy). Fron-
tiers In Ecology And Evolution, 7, 176.
Strani F., Bellucci L., Iannucci A., Iurino D.A., Mecozzi
B., Sardella R. (2021) - Palaeoenvironments of the
MIS 15 site of Cava di Breccia - Casal Selce 2
(central Italian Peninsula) and niche occupation of
fossil ungulates during Middle Pleistocene intergla-
cials. Historical Biology, in press.
Doi: 10.1080/08912963.2021.1935920
Terlato G., Bocherens H., Romandini M., Nannini N.,
Hobson K.A., Peresani M. (2018) - Chronological
and Isotopic data support a revision for the timing
of cave bear extinction in Mediterranean Europe.
Historical Biology, 31(4), 474-484.
Torres P. (1984) - Ursidos del pleistoceno-holoceno de
la Península Ibérica. Doctoral Dissertation,
E.T.S.I. Minas (UPM) pp. 653.
Torres P. (1988) - Osos (Mammalia, Carnivora, Ursidae)
del Pleistoceno Ibérico (U. deningeri Von Reiche-
nau, U. spelaeus Rosenmüller-Heinroth, U. arctos
Linneo). Boletín Geológico y Minero, 99, 4-940.
Tozzi C., Falgueres C., Mallegni F., Masini F., Negrino
F. (2000) - Les industries du Paléolithique inférieur
de Visogliano, Trieste, Italie, dans leur contexte
stratigraphique, biostratigraphique et géochronolo-
Wagner J., Čermák S. (2012) - Revision of the early
Middle Pleistocene bears (Ursidae, Mammalia) of
Central Europe, with special respect to possible co
-occurrence of spelaeoid and arctoid lineages.
Bulletin of Geosciences, 87(3), 461-496.
Wagner J., Jiangzuo Q., Lenardić J.M., Liu J. (2017) -
Taxonomic revision of bears from the locality Šan-
dalja I (Croatia) and its biostratigraphic conse-
quences. Fossil Imprint (Formerly Acta Musei Na-
tionalis Pragae, Series B-Historia Naturalis), 73(3-
4), 533-544.
gique. Les prémiers habitants de l'Europe, Tauta-
vel 10-15 avril 2000, Resumen de communica-
tions, pp. 104.
Valdiosera C.E., García N., Anderung C., Dalén L.,
Crégut-Bonnoure E., Kahlke R.D., Stiller M.,
Brandström M., Thomas M.G., Arsuaga J.L.,
Götherström A., Barnes I. (2007) - Staying out in
the cold, glacial refugia and mitochondrial DNA
phylogeography. Molecular Ecology, 16, 5140-
van Heteren A.H., Arlegi M., Santos E., Arsuaga J.L.,
Gómez-Olivencia A. (2019) - Cranial and mandib-
ular morphology of Middle Pleistocene cave bears
(Ursus deningeri): implications for diet and evolu-
tion. Historical Biology, 31(4), 485-499.
Wagner J., Sabol M. (2007) - Remarks on Biharian
bears (Ursidae: Ursus) from the territory of Slo-
vakia. Scripta Facultatis Scientiarum Naturalium
Universitatis Masarykianae Brunensis, 35, 159-
Review of Ursus material from Fontana Ranuccio (Middle Pleistocene, Central Italy)
Ms. received: November 17, 2020 Revised: March 23, 2021
Accepted: June 3, 2021 Available online: July 8, 2021
Conti J. et al.
... Other taxa include: Sus scrofa, Ursus deningeri, Ursus sp., Panthera sp., Crocuta crocuta, Canis mosbachensis, Macaca sylvanus and Homo sp. (Segre and Ascenzi 1984;Rubini et al. 2014;Conti et al. 2021). Small mammals are represented by few remains belonging to Microtus (Terricola) sp., Eliomys sp., cf. ...
The Ponte Galeria area within the city of Rome has yielded numerous fossiliferous localities that represent a reference point for the study of the European Middle Pleistocene ecosystems. Within Ponte Galeria a rich collection of fossil mammals has been unearthed from Cava di Breccia – Casal Selce 2 (MIS 15) thus the site represents an optimal laboratory to investigate the palaeoenvironments of a defined territory during the Middle Pleistocene. We investigate the feeding behaviours of the ungulate community of Cava di Breccia – Casal Selce 2 to reconstruct the MIS 15 habitats and also compare the data with those of the nearby site of Fontana Ranuccio (MIS 11) which shares similar faunal composition with Cava di Breccia – Casal Selce 2 to test if ungulates occupied the same niches during two different interglacials. Open habitats with scattered woodlands characterised the Ponte Galeria area during MIS 15, whereas woodlands were more widespread during MIS 11 at Fontana Ranuccio. Ungulates display similar diets in both localities, suggesting that cervids, large bovids and equids adopted the same niche partitioning strategies during both interglacials.
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The Middle Pleistocene was a crucial stage for the evolution of European mammals, a time when the majority of the modern taxa appeared in the continent for the first time. It is also in this interval that periodicity and intensity of glacial-interglacial cycles changed, an event that strongly impacted on Mediterranean marine and terrestrial ecosystems, and on vertebrate communities. This area can thus be considered an important laboratory to investigate how major climatic events influenced mammals' communities (among which also hominin populations) and the habitats they occupied. The state of art of the Middle Pleistocene palaeontological, archaeological and palaeoanthropological record of north Mediterranean region, and of the Italian Peninsula in particular, is here discussed.
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In this work, the Middle Pleistocene mammal assemblage from Ponte Molle, a historical locality of the urban area of Rome, has been revised together with a review of the stratigraphical succession of the deposit. This allows us to reconstruct the provenance of the fossil material and to provide chronological constrains trough the correlation with the lithostatigraphic and syn-themic units of the national geological cartography and the geochronologically-constrained aggradational units of the Paleo-Tiber reported in literature. The paleontological study together with the geological and stratigraphical review allow us to redefine the Ponte Molle deposit and its Middle Pleistocene faunal assemblage. In its new look, the age of the faunal assemblage from Ponte Molle could be referred to a time span ranging from 550 ka to 450 ka.
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After more than sixty years since its discovery, the fossils from Monte Peglia (late early Biharian, Umbria) are reconsidered in their entirety. The small mammals of Monte Peglia upper and lower levels had been studied in the past, whereas the remains of large mammals of Monte Peglia lower level, stored in several Institutions of central Italy, are described here for the first time. The following taxa have been described: Hystrix refossa, Homotherium latidens, Panthera cf. P. gombaszoegensis, Felis cf. F. lunensis, Canis mosbachensis, Vulpes alopecoides, Meles meles, Pannonictis cf. P. nestii, Mustela palerminea, Ursus cf. U. etruscus; Macaca sylvanus florentinus, Equus altidens, Stephanorhinus cf. S. hundsheimensis, Sus sp., Capreolus sp., Axis eurygonos, Hemitragus cf. H. orientalis, Bison degiulii. Moreover, the list of small mammals of the lower level has been updated with the addition of three new small vertebrate taxa: Rana sp., Myotis sp. (large size), cf. Miniopterus sp. The study of the remains of large mammals of the lower layer indicate the survival of a number of taxa of latest Villafranchian age. If we accept the biochronological correlation of Monte Peglia with the Colle Curti local fauna, its age should be ~1.072 Ma. In this case, it is possible to pinpoint the accumulation of the lower level to the MIS 35/33, as the small mammals confirm the presence of a mixed environment with forested and open spaces and warm temperate climate. The accumulation of the upper level, characterized by taxa typical of open spaces and steppes and a cooler climate, probably occurred during MIS 34/32.
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The techno-functional approach has been employed to better understand one of the more relevant artifact types generally found in Lower Palaeolithic sites: so-called small tools. Particularly, some Italian sites, such as Ficoncella, Isernia and others, have been the subject of specialized studies which provide evidence of an unexpected complexity of technical behaviours mainly related to highly specialized functional properties of the small tools. In this paper, we aim to enhance the debate on the topic by presenting a techno-functional study of the entire lithic assemblage coming from one of the most renown Middle Pleistocene sites in southern Europe, the open-air site of Fontana Ranuccio (Central Italy). Five groups of retouched tools have been identified: cutting tools, where retouch is usually applied to isolate a cutting edge on the blank; pointed tools, where retouch isolates a pointed edge; scrapers; and few other types of retouched tools such as notches and denticulates. We discuss a reconstruction of the reduction sequence in association with the functional features of the produced stone tools in order to better understand these Middle Pleistocene hominin behaviours. Broadly speaking, retouch seems to be used as a real technical process, not distinguishable from the reduction sequence. What seems relevant here is the need to modify the original morphology of flakes and cores in order to shape them into the final objectives of the production. In this perspective, blank production (débitage) and tool shaping (façonnage) are tightly interconnected one on the other.
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Following a recent chronostratigraphic revision of 17 fossiliferous sites hosting assemblages constituting local faunas of the Aurelian Mammal Age for peninsular Italy, we provide a re-structured biochronological framework and discuss the current validity and significance of the middle Pleistocene Faunal Units (FU) for this region. Contrasting with the previous model of a wide faunal renewal during Marine Isotope Stage (MIS) 9 (∼330 ka), the First Occurrences (FO) of several species of the Torre in Pietra FU are significantly backdated and referred to the Fontana Ranuccio FU (530-400 ka). We show that the faunal renewal was more gradual and occurred earlier than previously assumed. Many taxa that are typical of the late Pleistocene register their FO in the Fontana Ranuccio FU, latest Galerian, which is characterized by the almost total disappearance of Villafranchian taxa and by the persistence of typical Galerian taxa such as Dama clactoniana, Bison schoetensacki and Ursus deningeri, and by the FO of Stephanorhinus kirchbergensis, S. hemitoechus, Hippopotamus amphibius, Cervus elaphus eostepahnoceros, Ursus spelaeus, Canis lupus, and Vulpes vulpes. The next Torre in Pietra FU is characterized only by the FO of Megaloceros giganteus and Mustela putorius. However, we observe that MIS 9 marks the actual moment when the faunal assemblages of this region are represented only by those taxa characterizing the late middle Pleistocene and late Pleistocene. For this reason, we propose to still consider the Torre in Pietra (lower levels) local fauna as a conventional boundary for the Galerian-Aurelian transition. Finally, we remark that the strong faunal renewal in MIS 13, with five FOs, coincides with the temperate climatic conditions due to the absence of marked glacial periods that could have favored the FO and the subsequent spread of these taxa.
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Deninger’s bears (Ursus deningeri) have been studied less frequently than Ursus spelaeus s.l. Our objective is to present, for the first time, an analysis of the skull shape of U. deningeri. Bear crania and mandibles were digitised with a Microscribe or CT-scanned and the surface models subsequently landmarked. The landmarks were chosen based on a compromise between functional morphology and sample size. Results show that U. deningeri and U. spelaeus mandibles display very similar morphologies and allometric trajectories, both to each other and to Ailuropoda melanoleuca. It is inferred that masticatory adaptations to a herbivorous diet were already present in the Middle Pleistocene. U. deningeri displays a cranial morphology that is similar to that of U. spelaeus when comparing all species, but U. deningeri has a relatively narrower and dorsoventrally lower zygomatic arch than U. spelaeus, although the masticatory signal is less strong in the skull. We observe intraspecific differences between different populations of U. deningeri, which could parallel the genetic diversity found in U. spelaeus. The intraspecific differences found within U. deningeri may be temporal and/or geographical in nature and could be related to the evolution of the Late Pleistocene cave bear, but this hypothesis remains to be tested.
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Mastication of dietary items with different mechanical properties leaves distinctive microscopic marks on the surface of tooth enamel. The inspection of such marks (dental microwear analysis) is informative about the dietary habitus in fossil as well as in modern species. Dental microwear analysis relies on the morphology, abundance, direction, and distribution of these microscopic marks. We present a new freely available software implementation, MicroWeaR, that, compared to traditional dental microwear tools, allows more rapid, observer error free, and inexpensive quantification and classification of all the microscopic marks (also including for the first time different subtypes of scars). Classification parameters and graphical rendering of the output are fully settable by the user. MicroWeaR includes functions to (a) sample the marks, (b) classify features into categories as pits or scratches and then into their respective subcategories (large pits, coarse scratches, etc.), (c) generate an output table with summary information, and (d) obtain a visual surface‐map where marks are highlighted. We provide a tutorial to reproduce the steps required to perform microwear analysis and to test tool functionalities. Then, we present two case studies to illustrate how MicroWeaR works. The first regards a Miocene great ape obtained from through environmental scanning electron microscope, and other a Pleistocene cervid acquired by a stereomicroscope. We provide a new free and open‐access software (MicroWeaR) for dental microwear analysis. The tool allows for a rapid and inexpensive quantification and classification of the microscopic marks from a 2D image. Classification parameters and output settings are fully settable by the user.
The Ponte Galeria area within the city of Rome has yielded numerous fossiliferous localities that represent a reference point for the study of the European Middle Pleistocene ecosystems. Within Ponte Galeria a rich collection of fossil mammals has been unearthed from Cava di Breccia – Casal Selce 2 (MIS 15) thus the site represents an optimal laboratory to investigate the palaeoenvironments of a defined territory during the Middle Pleistocene. We investigate the feeding behaviours of the ungulate community of Cava di Breccia – Casal Selce 2 to reconstruct the MIS 15 habitats and also compare the data with those of the nearby site of Fontana Ranuccio (MIS 11) which shares similar faunal composition with Cava di Breccia – Casal Selce 2 to test if ungulates occupied the same niches during two different interglacials. Open habitats with scattered woodlands characterised the Ponte Galeria area during MIS 15, whereas woodlands were more widespread during MIS 11 at Fontana Ranuccio. Ungulates display similar diets in both localities, suggesting that cervids, large bovids and equids adopted the same niche partitioning strategies during both interglacials.
The scanty small mammals material of the Middle Pleistocene Fontana Ranuccio site is characterized by the presence of six species belonging to three order: Eulipotyphla, Lagomorpha, and Rodentia. The small mammal fossils evidence from Fontana Ranuccio, instead the scarce number of specimens, indicates that the site is characterized by a warm and wooded environment with subordinate grassland.
The Early and Middle Pleistocene were characterised by two major climatic events: the onset of the Quaternary glaciations and the Early-Middle Pleistocene Transition (EMPT) with glacial cycles changing their periodicity. The present study reviews recent research on the palaeoecological adaptations of Pleistocene ungulates from Mediterranean Europe following these global climatic changes, through an examination of dental wear patterns and hypsodonty indices of 27 fossil taxa from 4 key localities. Ungulates from Coste San Giacomo (2.1 Ma) adopted a wide range of feeding behaviours suggesting the presence of heterogeneous environments in the region. Following the gradual deterioration of climatic conditions and drier habitats ungulates display narrower diet ranges with no incidence of obligated browsers as attested at Olivola (~1.8 Ma). The diffusion of open habitats from the Early Pleistocene onwards is also reflected by the reduction of brachydont taxa. Discrepancy between dental mesowear and microwear patterns of Vallparadís Estació (~1.0–0.6 Ma) and Fontana Ranuccio (0.4 Ma) ungulates points to an increase in seasonality during the EMPT and after its consolidation. All examined taxa display dietary plasticity through the Early and Middle Pleistocene with even specialised groups, such as equids, showing a certain flexibility in their diet.