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A river otter hemimandible has been rediscovered during the revision of the historical collections of G.A. Blanc from Grotta Romanelli, complementing the ongoing multidisciplinary research fieldwork on the site. The specimen, recovered from the level G (“terre rosse”; early Late Pleistocene or late Middle Pleistocene), is here assigned to Lutra lutra . Indeed, morphological and morphometric comparisons with other Quaternary Lutrinae fossils from Europe allow to exclude an attribution to the relatively widespread and older Lutra simplicidens , characterized by distinctive carnassial proportions. Differences with Cyrnaonyx antiqua , which possessed a more robust, shellfish-feeding dentition, support the view of a successful niche repartition between the two species during the late Middle to Late Pleistocene of Europe. The occurrence of Lutra lutra from the “terre rosse” of Grotta Romanelli suggests deep modifications of the landscapes due to the ecological adaptation of the taxon, and indicates that the Eurasian otter spread into Europe at the Middle–Late Pleistocene transition.
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Rediscovering Lutra lutra fromGrotta Romanelli (southern Italy)
intheframework ofthepuzzling evolutionary history ofEurasian otter
BeniaminoMecozzi1 · AlessioIannucci1 · FabioBona2 · IlariaMazzini3 · PierluigiPieruccini4 ·
Received: 25 September 2020 / Accepted: 9 February 2021
© The Author(s) 2021
A river otter hemimandible has been rediscovered during the revision of the historical collections of G.A. Blanc from Grotta
Romanelli, complementing the ongoing multidisciplinary research fieldwork on the site. The specimen, recovered from the
level G (“terre rosse”; early Late Pleistocene or late Middle Pleistocene), is here assigned to Lutra lutra. Indeed, morpho-
logical and morphometric comparisons with other Quaternary Lutrinae fossils from Europe allow to exclude an attribution
to the relatively widespread and older Lutra simplicidens, characterized by distinctive carnassial proportions. Differences
with Cyrnaonyx antiqua, which possessed a more robust, shellfish-feeding dentition, support the view of a successful niche
repartition between the two species during the late Middle to Late Pleistocene of Europe. The occurrence of Lutra lutra from
the “terre rosse” of Grotta Romanelli suggests deep modifications of the landscapes due to the ecological adaptation of the
taxon, and indicates that the Eurasian otter spread into Europe at the Middle–Late Pleistocene transition.
Keywords Pleistocene· Mediterranean· Carnivora· Body mass· Lutrinae
Otters (Lutrinae), subfamily of Mustelidae, constitute a
widely distributed carnivoran group (13 extant species),
which evolved morphological adaptations for living and
hunting in aquatic environments (Wilson and Mittermeier
2009; Nowak 1999). Despite earlier insights from morpho-
logical analysis (Van Zyll de Jong 1972,1987), river otters
have been traditionally ascribed only to Lutra, until genetic
data highlighted the substantial molecular divergence
between the Eurasian and American branches of Lutrinae
evolutionary tree, now assigned to Lutra and Lontra, respec-
tively (Koepfli and Wayne 1998; Koepfli etal. 2008).
However, if Lontra occurred in North American depos-
its since the Pliocene (Prassack 2016), placing the first
appearance of Lutra in a chronological framework is on the
contrary complicated by the use of the genus either as a
waste-basket taxon, or in sensu lato, to allocate specimens
of uncertain affinities and/or Lutra-like morphology (Wil-
lemsen 1992; Geraads etal. 2015; Cherin etal. 2016). In
Handling Editor: Irina Ruf.
* Beniamino Mecozzi
Alessio Iannucci
Fabio Bona
Ilaria Mazzini
Pierluigi Pieruccini
Raffaele Sardella
1 PaleoFactory Laboratory, Department ofEarth Sciences,
Sapienza University ofRome, P.le Aldo Moro 5,
00185Rome, Italy
2 Museo Civico dei Fossili di Besano, Via Prestini 5,
21050Besano, VA, Italy
3 Institute ofEnvironmental Geology andGeoengineering
(IGAG), CNR, Area della Ricerca di Roma 1, Montelibretti,
Via Salaria Km 29,300, Monterotondo, 00015Rome, Italy
4 Department ofEarth Sciences, University ofTorino, Via
Valperga Caluso 35, 10125Turin, Italy
B. Mecozzi etal.
1 3
fact, the earliest “Lutra” species, Lutra affinis Gervais 1859,
is only known from three sites close to the Miocene–Plio-
cene boundary: Venta del Moro (Spain), Maramena (Greece)
and Montpellier (France). However, the whereabouts of the
never-figured type material from Montpellier are unknown
(Willemsen 1992). Furthermore, the fragmentary remains
from the other localities markedly differ in size (Koufos
2011; Montoya etal. 2011) and have been suggested to rep-
resent different taxa (Cherin etal. 2016). Several putative
Lutra species have been described from the Pliocene to the
Early Pleistocene of Europe (Gervais 1859; Pohle 1919;
Kurtén 1968; Willemsen 1992; Fejfar and Sabol 2004), but
none of them provides conclusive evidence of their generic
status (Cherin etal. 2016).
A long gap characterized the Lutrinae record during the
late Pliocene and the beginning of the Early Pleistocene,
until the earliest occurrence of Lutra simplicidens and
Lutraeximia umbra from the late Villafranchian Upper Val-
darno (Tuscany) and Pantalla (Umbria), respectively (Cherin
and Rook 2014; Cherin etal. 2016). The latter taxon is only
known from the holotype cranium, whereas the former is
relatively widespread in the early Middle Pleistocene and
is regarded as ancestral to several endemic otters of the
Mediterranean islands (Willemsen 2006): “Lutra” euxena
from Malta, Lutraeximia trinacriae from Sicily, and Sar-
dolutra ichnusae, “Lutra” castiglionis, Algarolutra majori
and Megalenhydris barbaricina from Sardinia and Corsica,
even though these fossil samples often include few and
fragmentary remains (Willemsen 1992; Cherin etal. 2016).
Nonetheless, the extant Eurasian river otter, Lutra lutra,
is considered not related to these European species and its
origin is placed in the Asian continent, probably deriving
from the fossil species Lutra palaeoindica (Willemsen 1992;
Raghavan etal. 2007). This scenario has received indirect
support from the first description of L. simplicidens max-
illary dentition, which led Cherin (2017) to doubt of the
species generic attribution, remarking the similarities with
Lutraeximia umbra (Cherin etal. 2016). Alternatively, Fejfar
and Sabol (2004) argued for a derivation of the extant Lutra
sumatrana from Lutra bravardi Pomel 1843, but excluding
a close relationship with L. lutra of the latter species, which
is difficult to reconciliate with molecular data that consist-
ently find a close relation between L. lutra and L. sumatrana
(Koepfli etal. 2008; Hung and Law 2016).
L. lutra is the most widely distributed otter species, with
extant populations occurring throughout Asia, Europe and
northern Africa. Historically, this carnivoran was also wide-
spread in Japan, Portugal, arctic region of Asia and Europe
and southern Indonesia (Foster-Turley etal. 1990). Up to
now, fossils of the species are mainly known from Holocene
deposits (Sommer and Benecke 2004), even if it was also
reported from the late Middle Pleistocene of Grotta Mag-
giore di San Bernardino (Cassoli and Tagliacozzo 1994c)
and the Late Pleistocene sites of Geographical Society
Cave (Russian Far East) (Baryshnikov 2015), Pilisszánto
1 (Jánossy 1986) and Skjonghelleren Cave (Larsen etal.
1987). Indeed, Baryshnikov and Puzachenko (2012) sug-
gested that the river otter could have survived during the
cold phases of the Last Glacial in southern European refu-
gia, from where it redispersed during the Early Holocene.
Another otter species, Cyrnaonyx antiqua, is reported from
several late Middle to Late Pleistocene European localities
and is characterized by a more robust, shellfish-feeding type
of dentition (Kurtén 1968; Willemsen 1992).
The otter fossil from Grotta Romanelli (southern Italy),
belonging to the historical collection of Gian Alberto Blanc
and never figured or described, is here presented for the first
time. In addition, morphological and morphometric com-
parisons with other Lutrinae fossils coming from the Early
Pleistocene to Middle Holocene sites of Europe and with
extant specimens of European otter are taken into account
to evaluate their morphological and body size variability.
Materials andmethods
Grotta Romanelli (40° 00 58.33 N, 18° 25 59.36 E) is
a costal cave located in the administrative territory of the
Castro (Lecce) municipality, at the south-eastern extremity
of Apulia (southern Italy) (Fig.1a–d). The cave lies within
the area of the Regional Natural Park “Costa Otranto—Santa
Maria di Leuca and Bosco di Tricase”. From the beginning
of the twentieth century, Grotta Romanelli was considered
a key site for Mediterranean Quaternary studies, due to its
remarkable importance as the first evidence of the Upper
Paleolithic period in the Italian Peninsula and the first record
of rock art in the Euro-Mediterranean area (Graziosi 1973;
Sardella etal. 2018, 2019). The deposit was discovered in
1874 by Ulderigo Botti (Botti 1874), but only during the
earlier 1900s, Paolo Emilio Stasi realized the first excavation
campaigns (Stasi and Regalìa 1904). In 1914, Gian Alberto
Blanc began a pioneering excavation campaign, during
which the first systematic archeological/paleontological
and stratigraphic study using scientific methods was car-
ried out (Blanc 1920, 1928) (Fig.1e–g). Nowadays, Grotta
Romanelli is recognized as a key site for the Mediterranean
Pleistocene for its archeological and paleontological content
and the relative stratigraphical, geomorphological and radio-
metric data (Sardella etal. 2018, 2019).
The infilling deposit was described by Blanc (1920)
which identified two main stratigraphical complexes: the
upper complex, called “terre brune” (levels A–E), bearing
Upper Paleolithic artifacts and vertebrate fauna; the lower
complex (G–K) formed by a deposit called “terre rosse”
(level G), bearing Middle Paleolithic artifacts and verte-
brate fauna, a stalagmitic layer (level H), a bone breccia
Lutra lutra from Grotta Romanelli and the puzzling evolutionary history of Eurasian otter
1 3
(level I), and a beach deposit (level K) referred to the Tyr-
rhenian Stage (Marine Isotopic Stage 5) (Blanc 1920, 1928)
(Fig.1f, g). These two complexes were separated by a
thick stalagmitic layer (called level F) (Blanc 1920, 1928)
(Fig.1g). The upper complex was referred to the end of
the Late Pleistocene and Early Holocene on the basis of
radiocarbon datings performed since the end of 1950
(Bella etal. 1958; Vogel and Waterbolk 1963; Alessio etal.
1964, 1965). The two speleothems (levels H and F) were
also dated with the 230Th/234Th method: level H yielded an
apparent age of < 69,000years BP, and level F was dated to
40,000 ± 3250years BP (Fornaca-Rinaldi 1968a, b; Fornaca-
Rinaldi and Radmilli 1968; Sardella etal. 2018). During the
1950s and 1960s, the excavation campaigns were directed
by Cardini with the support of the Italian Institute of Human
Paleontology (IsIPU) (Sardella etal. 2018).
In 2015, new multidisciplinary research fieldwork began,
coordinated by Sapienza University of Rome together with
other research institutions and with the authorization of
Soprintendenza Archeologia Belle Arti e Paesaggio di
Brindisi, Lecce e Taranto, with the aim to provide an updated
stratigraphic and chronological scheme as well as to describe
the fossil remains and artifacts (Giustini etal. 2018; Sardella
etal. 2018, 2019), which is also triggering a revision of the
large mammal faunas from the region (Sardella etal. 2014;
Mecozzi etal. 2017, 2018, 2019; Iannucci etal. 2020; Iurino
Fig. 1 Geographical position of Grotta Romanelli (ac). External
(d) and internal (e) views of the cave. Stratigraphy of the deposit as
described by Blanc (f), the level where the fossil otter was found is
highlighted in light orange. Schematic log of the sedimentary suc-
cession (modified by Blanc 1920). Fossil Lutra lutra from Grotta
Romanelli in labial (h), lingual (i) and occlusal (j) views
B. Mecozzi etal.
1 3
etal. 2020). The research team had to face the consequences
of more than 40years of inactivity in the site and the com-
bined effects of erosion and illegal excavations that took
place during last decades (Conti etal. 2017; Giustini etal.
2018; Sardella etal. 2018). Recently, the age of the “terre
brune” has been reassessed, with new fossil remains submit-
ted for 14C dating using the accelerator mass spectrometry
radiocarbon (AMS14C) dating techniques (Calcagnile etal.
2019; Sigari etal. 2021). The results expand and refine the
previous chronology, with a time span for levels D and B,
respectively, from 13,976–13,545 14C cal BP to 9135–8639
14C cal BP (Calcagnile etal. 2019), whereas the level E was
dated between 13,087–12,839 and 13,408–13,153 (Sigari
etal. 2021). As reported by Sardella etal. (2018), one of the
main unexplained aspects, which needs to be investigated,
concerns the age of the lower deposits due to the apparent
contradiction between the supposed archaic nature of the
Mousterian artifacts and the position of the vertebrate fauna
being situated above the supposed MIS 5.5 terrace.
New analyses on the fauna and lithic tools from the lower
complex suggest a possible late Middle Pleistocene to early
Late Pleistocene age (Sardella etal. 2018), but more data
need to define the updated chronostratigraphy of the infill-
ing deposit.
The fossil remains described in this work were collected
from the “terre rosse” (level G), but no information about the
exact stratigraphical position is available (Blanc 1920, 1928)
(Fig.1h–j). Other mammal species found in the level G are:
Elephas antiquus (= Palaeoloxodon antiquus), Rhinoceros
merckii (= Stephanorhinus kirchbergensis), Hippopota-
mus sp. amphibius, Hippopotamus sp. pentlandi, Bos tau-
rus var. primigenius (= Bos primigenius), Cervus elaphus,
Cervus elaphus var. corsicanus (= Cervus elaphus corsi-
canus), Dama dama, Capreolus capreolus, Equus caballus,
Hyaena crocuta var. spelaea (= Crocuta crocuta spelaea),
Canis lupus, Canis aureus, Meles taxus (= Meles meles),
Oryctolagus cuniculus and Pelagius monachus (= Pelagius
monachus) (Blanc 1920). The wolf and rhino remains from
level G have recently been revised by Sardella etal. (2014)
and Pandolfi etal. (2017), who referred them to Canis lupus
and Stephanorhinus hemitoechus, respectively.
The fossil material from Grotta Romanelli is housed in
the collection of the “Museo delle Civiltà” (ex. Museo Preis-
torico ed etnografico Luigi Pigorini), labeled with a progres-
sive number MUCIV-GR1935 (Fig.1h–j).
According to von den Driesch (1976) and Willemsen
(1992), 16 mandibular and dental variables have been con-
sidered: total length (Tl), length from the anterior border of
the C1 to posterior border of the M2 (LC1M2), height of the
mandibular corpus above M1 (HM1), length and breadth of
the lower teeth (C1L, C1W, P2L, P2W, P3L, P3W, P4L, P4W,
M1L, M2L, M2W) length of trigonid (M1Ltri), breadth of
the trigonid (M1Wtri) and talonid (M1Wtal). Craniodental
morphometric measurements were recorded to the nearest
0.1mm with a digital caliper.
Literature data on fossil river otters from the Early Pleis-
tocene to Early Holocene (Late Villafranchian to Aurelian)
sites (Table1) as well as extant specimens of L. lutra have
been used. Comparisons are also based on fossil material
found in the late Neolithic site of Tosina di Monzambano
(Mantova—Italy) (Bona 2014; Poggiani Keller 2014) and
extant specimens housed in the PaleoFactory laboratory,
Department of Earth Sciences, Sapienza, University of
We performed body mass estimates of the fossil sample
following regression equations based on the length of the
lower carnassial (Van Valkenburg 1990) (Table1). Size and
proportions of the lower carnassial of the specimen from
Grotta Romanelli were compared with literature data on
Pleistocene–Holocene otters through standard bivariate plot.
In mustelids, these teeth enable determining masticatory
performance and bite force and their morphology and size
are linked to diet (e.g., Baryshnikov etal. 2003; Madurell-
Malapeira etal. 2011a, b).
Systematic Paleontology
ClassMammalia Linnaeus, 1758
OrderCarnivora Bowdich, 1821
FamilyMustelidae Fischer von Waldheim, 1817
SubfamilyLutrinae Bonaparte, 1838
GenusLutra Brisson, 1762
Lutra lutra Linnaeus, 1758
Material. Right hemimandible (MUCIV-GR1935; measure-
ments in Table2).
Locality and level. Grotta Romanelli; “terre rosse”—G.
Description. MUCIV-GR1935 is a finely preserved mandible,
lacking only of a portion of the vertical ramus. In labial view,
it shows three mental foramina: one located to the mid length
of the C1; one posteriorly to the distal border of the P2; and
one to the mid length of the P3. The anterior portion of the
masseteric fossa reaches the level of the distal portion of the
M1. The ramus is high and straight with respect to the corpus
axis and the angular process is elongated dorsally. In dorsal
view, the mandibular condyle has a straight posterior border
and is inclined in posterolateral direction. The ventral outline
of the mandibular corpus is almost straight, but anteriorly, at
the level of the alveolus of the P2, it sharply rears upwards. The
toothrow is relatively straight and the longitudinal axis of each
Lutra lutra from Grotta Romanelli and the puzzling evolutionary history of Eurasian otter
1 3
tooth diverges, with the C1, P3 and M1 axes almost aligned.
The posterior part of the P4 crown slightly overlaps with the
anterior margin of the M1. No incisors are preserved. The C1
is mesodistally elongated, with a smooth crown. No diastema
separates the C1 from the P2. The P2 is a two-rooted and buc-
colingually compressed tooth with a small and mesially pro-
jected protoconid, and with a crest connecting the weak distal
cingulum to the protoconid. The P3 is buccolingually com-
pressed and has the high, straight and anteriorly located pro-
toconid, and a weak crest connecting the mesial cingulum to
the protoconid. It has weak lingual and labial cingula. The P4
is also buccolingually compressed, with the high protoconid,
a marked distal accessory cusp, and a marked basal cingulum.
In the lower carnassial (M1) the talonid is wider than
the trigonid. The paraconid is well developed and high
as the metaconid. The protoconid is the highest cusp and
two distinct grooves separate it from the paraconid and
the entoconid. The entoconid and hypoconid are reduced
and aligned with the metaconid and the paraconid, respec-
tively. Along the distal rim of the talonid, there are two
very reduced cusps, buccally the entoconulid and lingually
the hypoconulid. The talonid basin is concave. The basal
distal cingulum is well developed, whereas anteriorly
tends to be less marked.
The M2 displays a circular outline, with four cusps. The
less developed cuspids, the paraconid and the hypoconid,
occupy the anterior and posterior portion of the toot,h
respectively; whereas, the metaconid and the protoconid
are more developed.
Table 1 Literature data on fossil otters from the Early Pleistocene to Holocene sites of Eurasia
Species Site Catalog number Age References M1L (mm) Body size (kg)
Lutra lutra Grotta Romanelli MUCIV-GR1935 late Middle–early
Late Pleistocene
This work 12.8 6.5
Lutra lutra Primorskii Territory ZIN 37285–1 Late Pleistocene Baryshnikov (2015) 13.7 8.2
Lutra lutra Ellewoutsdijk RGM 146662 Holocene Willemsen (1992) 12.4 5.8
Lutra lutra Kapel-Avezaath RGM 147002 Holocene Willemsen (1992) 13.4 7.6
Lutra lutra Maasvlakte Ke RM238 Holocene Willemsen (1992) 13.3 7.4
Lutra lutra Maasvlakte Ke RM2974 Holocene Willemsen (1992) 12.5 6.0
Lutra lutra Maasvlakte Ke RM 4109 Holocene Willemsen (1992) 13.6 8.0
Lutra lutra Weesp IPP We2/1 Holocene Willemsen (1992) 13.2 7.2
Lutra lutra Vlaardingen IPP G20d/B7 Holocene Willemsen (1992) 13.0 6.9
Lutra lutra Vlaardingen IPP 121c/B4 Holocene Willemsen (1992) 12.6 6.2
Lutra lutra Vlaardingen IPP F20b/B2 Holocene Willemsen (1992) 13.0 6.9
Lutra lutra Leidschendam IPPL387/8 Holocene Willemsen (1992) 11.9 5.0
Lutra lutra Tosina Tos 6813 Holocene This work 13.6 8.0
Lutra lutra Tosina Tos 7128 Holocene This work 13.1 7.0
Lutra simplicidens Chumbur Kosa AMZ/OP-966 Early Pleistocene Sotnikova and Titov (2009) 13.1 7.0
Lutra simplicidens Upper Valdarno NHM M296688 Early Pleistocene Cherin and Rook (2014) 12.6 6.2
Lutra simplicidens Breccia di Soave Na Middle Pleistocene Pasa (1947) 12.0 5.2
Lutra simplicidens Hundsheim PIV Hsh VII/26 Middle Pleistocene Thenius (1965) 12.8 6.5
Lutra simplicidens East Runton BM 6089 Middle Pleistocene Willemsen (1992) 12.0 5.2
Lutra simplicidens West Runton CM CR4 984 Middle Pleistocene Willemsen (1992) 12.7 6.3
Lutra simplicidens Mosbach SMF PA/F8718 Middle Pleistocene Willemsen (1992) 12.9 6.7
Lutra simplicidens Voigtstedt KMBF I-711K Middle Pleistocene Willemsen (1992) 12.6 6.2
Cyrnaonyx antiqua Lunel Viel LPM LUV 68 Middle Pleistocene Serres etal. (1839) 14.0 8.9
Cyrnaonyx antiqua Saalfeld MNB MBMa8069 Late Pleistocene Willemsen (1992) 14.0 8.9
Cyrnaonyx antiqua Ehringsdorf Na Late Pleistocene Willemsen (1992) 14.0 8.9
Cyrnaonyx antiqua Maasvlakte Ke RM 226 Late Pleistocene Willemsen (1992) 14.8 10.8
Cyrnaonyx antiqua Tornetown Cave BM M 34371 Late Pleistocene Willemsen (1992) 13.5 7.8
Cyrnaonyx antiqua Tornetown Cave BM M 34372 Late Pleistocene Willemsen (1992) 13.2 7.2
Cyrnaonyx antiqua Tornetown Cave BM M 50708 Late Pleistocene Willemsen (1992) 14.1 9.1
Extant Lutra lutra Italian Peninsula 882 This work 13.4 7.6
Extant Lutra lutra Italian Peninsula 882 This work 13.5 7.8
Extant Lutra lutra Italian Peninsula 883 This work 13.8 8.4
B. Mecozzi etal.
1 3
Comparison. MUCIV-GR1935 possesses a typical Lutra-
like appearance. Morphologically, the lower carnassial of
the specimen from Grotta Romanelli shares several features
with L. lutra (Fig.2f–l): talonid wider than trigonid; con-
cave talonid; hypoconulid present; strong and continuous
lingual cingulum (Willemsen 1992). Opposite characters
were observed in L. simplicidens from Hundsheim, Sussen-
born, Mosbach, East Runton and West Runton (Fig.2a–d):
trigonid wider than talonid; flat talonid; hypoconulid absent;
less developed and incomplete lingual cingulum (Willem-
sen 1992). However, the specimen NHM M29688 from the
Upper Valdarno ascribed to L. simplicidens possesses a
continuous cingulum, which extends all around the paraco-
nid, and the trigonid and talonid are equal in width (Cherin
and Rook 2014) (Fig.2a). Similarly, in the specimen AMZ/
OP-966 from the Chumbur Kosa (Azov Sea area), classi-
fied as L. simplicidens tamanensis, trigonid and talonid are
equal in width (Sotnikova and Titov 2009) (Fig.2d). The
studied specimen differs from L. palaeindica from the Upper
Siwalik in that it possesses a larger trigonid than talonid
(Fig.2e). MUCIV-GR1935 shares several features with the
extant L. sumatrana (Fig.2m, n), such as the talonid larger
than trigonid and the concave talonid basin.
According to Willemens (1992), the lower carnassial of
Cyrnaonyx antiqua differs from L. lutra in the following fea-
tures: larger paraconid; metaconid more closely connected
to the other two cuspids of the trigonid and smaller than
the other two cuspids; talonid wider than trigonid; labial
cingulum more developed; larger hypoconid (Fig.2o, p).
MUCIV-GR1935 from Grotta Romanelli resembles C.
antiqua for the large paraconid and the talonid wider than
In the extant specimens of L. lutra, the morphology of the
M2 is highly variable, with three main morphotypes recog-
nized, characterized by a progressive complication of crown
morphology: A1, with only protoconid and metaconid; A2,
with protoconid, metaconid, paraconid and hypoconid;
A3, with all the four cuspids on the grinding surface and
an additional element, a projection on the lingual side of
tooth crown (Gimranov and Kosintsev 2012). Following this,
MUCIV-GR1935 can be referred to the A2 morphotype. The
only M2 of L. simplicidens from the Upper Valdarno differs
from MUCIV-GR1935 in the absence of the paraconid and
in possessing a more centrally placed protoconid (Cherin
and Rook 2014).
Morphometric comparison
Standard bivariate plots of talonid width (M1WTal) vs length
(M1L) of the lower carnassial show a distinctively small size
for the genus Lutra (Fig.3a). Within the Lutra group, the
specimen from Grotta Romanelli and those of L. simplici-
dens fall in the broader variability of L. lutra. The C. antiqua
material is instead characterized by a large size of the lower
carnassial, well distinct in the M1WTal (Fig.3a). Consider-
ing the M1WIndex (calculated as M1Wtal−M1Wtri), other
differences can be observed (Fig.3b). Two groups can be
distinct based on the M1WIndex: the material with positive
values (above the dotted line), which has a larger talonid;
and those specimens with negative values (below the dotted
line), which have a larger trigonid. Generally, C. antiqua
and L. lutra occupy the upper part of the graphs, whereas L.
simplicidens occupies the lower part (Fig.3b). However, a
few specimens of Lutra show a talonid and trigonid equal in
width (located on the dotted line) (Fig.3b).
Body mass
Body size in extant otters is hugely variable, reflecting
the diverse ecological niches this group occupies (Con-
roy etal. 2000; Kruuk 2006): small-sized piscivore (c.
5–10kg), medium-sized cancrivore (c. 11–12kg) and
large-sized molluscivore (c. 22–25kg) (Raghavan etal.
2007). Therefore, in the fossil record, reconstructing the
body size could represent an important proxy to evalu-
ate possible ecological niche differentiation (Conroy etal.
2000). The Eurasian otter is a dimorphic species, with
males 50% larger than females (Larivière and Jennings
Table 2 Mandibular and lower teeth measurements (mm) of Lutra
lutra from Grotta Romanelli
Measurements Grotta Romanelli
TL 66.5
C1L 8.0
C1W 4.9
P2L 4.8
P2W 3.1
P3L 5.4
P3W 3.5
P4L 7.6
P4W 4.5
M1L 12.8
M1Ltri 7.5
M1Wtri 5.7
M1Wtal 6.3
M2L 4.8
M2W 5.3
Lutra lutra from Grotta Romanelli and the puzzling evolutionary history of Eurasian otter
1 3
2009). According to Conroy etal. (2000), the body mass
of males is of 5.45–11.4kg, whereas that of females is
of 3.36–7.6kg. Conversely, values reported by Kruuk
(2006) range from 5.0 to 14.0kg, but no sex distinction
was made. Considering the fossil sample, the M1L pre-
dicts a reduced body-size variation for the Early–Middle
Pleistocene L. simplicidens, with values between 5.2 and
7.0kg (Table1). The L. lutra sample possesses slightly
larger, but not different body mass, with estimated values
ranging from 5.0 to 8.2kg (Table1). Finally, the highest
body size values are obtained for the sample of C. antiqua,
whose estimates are between 7.8 and 10.8kg (Table1).
The estimated values on fossil otters reveal a reduced
body size variation among groups, with slight difference in
the average values and wide overlap in distribution ranges.
Taking into account the body mass range of the extant
Eurasian otter, the difference among the fossil groups is
even less significant, since all their body size values fall
into the extant variation range.
Eurasian quaternary otters
Blanc (1920, 1928) reported for the first-time the presence
of L. lutra from the level G of Grotta Romanelli. Since then,
the Eurasian otter from this archeological site was cited by
paleontologists (e.g., Willemens 1992; Pennacchioni and
Cassola 1986), but no operational diagnosis was available,
thus preventing its morphological and/or biometric compari-
son. The lack of these data sometimes sparked debates on
taxonomy of some specimens belonging to these collections,
as for instance happened for the canid collection recovered
from the level G of Grotta Romanelli (see Sardella et al.
2014 for discussion). Morphological analysis reveals that the
features of the fossil specimen from Grotta Romanelli clearly
fit with an attribution to the extant L. lutra, confirming what
Blanc (1920, 1928) reported.
Fig. 2 Comparative morpholog-
ical sketch of the lower carnas-
sial of the fossil and extant otter
taxa in occlusal view (drawn not
in scale). Lutra simplicidens: a
NHM M29688 from Upper Val-
darno (Cherin and Rook 2014);
b BM 6089 from West Runton
(Willemsen 1992); c SMF PA/F
8718 from Mosbach (Willemsen
1992); d KP-27075/OP-966
from Chumbur Rosa (Sotnikova
and Titov 2009). Lutra palaein-
dica: e BM37152 from Upper
Siwalk (Matthew 1929). Fossil
Lutra lutra: f MUCIV-GR1935
from Grotta Romanelli; g ZIN
37,285–1 from Geographical
Society Cave (Baryshnikov
2015); h, i Tos 6813 and Tos
7128 from Tosina di Mon-
zambano (Mantova—Italy).
Extant Lutra lutra: j, k 882 and
883; l n/a (Pohle, 1919). Lutra
sumatrana: m n/a (Pohle 1919);
n ZM CN4494 (Zyll de Yong,
1972). Cyrnaonyx antiqua:
oq BM M 34,378 and BM M
50,709 from Tornetown Cave
(Willemsen 1992); p Ke RM
226 from Maasvlakte (Willem-
sen 1992)
B. Mecozzi etal.
1 3
Despite several fossil otters have been referred to Lutra
over the years, many of them are now reassigned to other
genera, whereas “Lutra” is retained mainly for ease of ref-
erence for taxa of uncertain taxonomy (Willemsen 1992;
Cherin etal. 2016), recognizing its paraphyletic nature (Ger-
aads etal. 2015). L. simplicidens has a relatively abundant
fossil record, compared to other Pleistocene Lutrinae. How-
ever, it is almost entirely known from isolated mandibular
and postcranial remains (Willemsen 1992), which are not
directly comparable with the recently described Lutraeximia
umbra, only known for the holotype cranium. It is worth
noticing that Cherin (2017), providing the first description
of the maxillary dentition of L. simplicidens, recognized a
close relationship between this species, Lutraeximia umbra,
Lutraeximia trinacriae and other Mediterranean Quaternary
otters likely forming a monophyletic clade (Cherin etal.
2016). If new discoveries are needed to substantiate the
attribution of L. simplicidens to Lutraeximia (Cherin 2017),
nonetheless there is general agreement in recognizing as
hardly conceivable a derivation of L. lutra from L. simplici-
dens (Willemsen 1992, 2006). Our data agree in evidencing
morphometric differences between the two species, provid-
ing further support for a different derivation of L. lutra.
Willemsen (1992) hypothesized an Asian origin for L.
lutra, probably from L. palaeindica. However, the latter
taxon is only known from the type sample (Falconer 1868;
Fig. 3 Plot of lower carnassial:
a length of M1 vs. width of
M1 talonid; b length of M1 vs.
width of M1 W index (calcu-
lated as M1 W talonid−M1 W
Lutra lutra from Grotta Romanelli and the puzzling evolutionary history of Eurasian otter
1 3
Pohle 1919; Mattthew 1929), and possibly from another
specimen (Raghavan etal. 2007), recovered from the Upper
Siwalik. Similarly, no cranial material of L. simplicidens
has been described so far, which highlights the strongly
restricted knowledge on the morphological variability of
these species. Nevertheless, in dental morphology, L. pal-
aeindica shares many features with L. simplicidens. In the
P4: the metacone lower than the paracone with a crest that
connects the two cusps curved lingually, the parastyle well
defined and the talonid mesiodistally elongated, triangular-
shaped in occlusal view (though more pronounced in L.
palaeindica) (Fig.4); and in the M1: the trigonid larger the
talonid and a relatively reduced metaconid (Fig.2). Com-
pared to C. antiqua, these two taxa instead possess marked
differences in the upper and lower carnassial morphology,
especially in the P4, where the profile in occlusal view is
trapezoidal (Fig.4j–m).
Setting our results in chronological framework, it seems
that L. lutra dispersed into Europe from Asia at the begin-
ning of the Late Pleistocene or, possibly, during the late
Middle Pleistocene, replacing L. simplicidens. In this sce-
nario, the taxonomic attribution of two Middle Pleistocene
specimens needs to be considered. The first is the hemi-
mandible from Cengelle BB classified as L. lutra by Pasa
(1947). According to his description, the specimen displayed
clear Lutra features, among which is a trigonid larger than
talonid in the lower carnassial. It is worth noting that L.
simplicidens was instituted by Thenius (1965), and thus the
name was not available to Pasa (1947). According to Wil-
lemsen (1992), the trigonid larger than talonid character has
a high taxonomic value, which allows an unquestionable
attribution of the specimen from Cengelle BB to L. simplici-
dens. The second specimen was a P4 recovered from Hoxne
and ascribed to Lutra sp. by Willemsen (1992). The author
recognized the strong resemblance with L. lutra, but given
the absence of coeval maxillary Lutrinae material, he could
not rule out an attribution to L. simplicidens, preferring an
evolutionary hypothesis in which L. lutra appears during the
Early Holocene. Recently, Cherin (2017) described for the
first time the morphology of the maxillary dentition of L.
simplicidens. The comparison of the P4 shows slightly but
significative morphological differences between the Hoxne
specimen and Voigtstedt ones (Fig.4a, b, d; Table3). Nota-
bly, L. simplicidens possesses a characteristic talon profile,
with a convexity along the distal margin, about half length,
in occlusal view (Cherin 2017) (Fig.4a, b). In the P4 from
Hoxne, the profile of the distal margin is straight, not con-
vex, which allows to refer it to L. lutra.
Paleoecological implications
Despite its fossil record is less abundant if compared to other
Mustelidae taxa (e.g., Meles meles, Mecozzi etal. 2019),
otters are largely distributed across the Eurasian continent
over a long stratigraphical interval. Currently, this carnivoran
inhabits a large variety of aquatic habitats, including high-
land and lowland lakes, rivers, streams, marshes, swamp for-
ests and coastal areas independent of their size, origin or lat-
itude (Mason and Macdonald 2009; Hung and Law 2016). It
can be found in brackish waters from sea level up to 4120m
(Mason and Macdonald 2009). Similarly to other European
carnivorans, the extant populations of L. lutra suffer the
Fig. 4 Upper fourth premolar of fossil and extant otter in occlusal
view (drawn not in scale). Lutra simplicidens: a IQW 1966/7313
(Voi. 3525) from Voigtstedt; b IQW 1966/7314 (Voi. 3532) from
Voigtstedt (Cherin 2017). Lutra palaeindica: c BM37151 from Upper
Siwalik (Matthew 1929). Lutra sp.: d BM4936 from Hoxne (Willem-
sen 1992). Lutra lutra: e n/a (Zyll de Jong 1972). f n/a (Pohle 1919).
Lutra sumatrana: g CN4494 (Zyll de Jong 1972); h n/a (Pohle 1919).
Lutraeximia umbra: i SBAU 337,654 from Pantalla (Cherin et al.
2016). Cyrnaonyx antiqua: j 7M from Cengelle (Pasa 1947); km
BM34371, BM50707 and BM34373 from Tornetown Cave (Willem-
sen 1992)
B. Mecozzi etal.
1 3
strong human pressure, which mainly reflects the habitat
loss (Toschi 1965; Prigioni 2003). In the Italian Peninsula,
the presence of the Eurasian otter is nowadays confined in
the southern regions, with stable populations reported also
in northern Apulia (Toschi 1965; Prigioni 2003). Thus, the
occurrence of L. lutra from Grotta Romanelli is surprisingly,
since the geographical position of the site falls outside the
range of the extant species. Moreover, its ecological profile
indicates a strong relation to large freshwater bodies (rivers
or lakes), which are no longer present in the southern part
of the Apulia (Mastronuzzi and Sansò 2017). Therefore, its
presence strongly supports a more diversified hydrographic
network in this territory during the late Quaternary.
Body mass estimates reveal a large overlap between L.
lutra and L. simplicidens (Fig.5), which also share a fish-
feeding type of dentition, making difficult to envision a pos-
sible coexistence. On the contrary, C. antiqua has similar but
slightly larger values of estimated body mass but occupied a
different trophic niche. In fact, given its robust dentition with
broader teeth, several authors aligned it with Aonyx (e.g.,
Malatesta 1978; Pennacchioni and Cassola 1986), whose
paleobiological interpretation points to a more important
role of shellfish in its diet (Willemsen 1992). Following
this, a sympatric coexistence scenario of the fish-feeding
L. simplicidens and L. lutra with C. antiqua could be pos-
sible, with different craniodental features which reflect the
Table 3 Literature data on
the upper fourth premolar of
fossil otters from the Early
Pleistocene to Holocene sites of
Species Site Specimen number References P4L (mm)
Lutraeximia umbra Pantalla SBAU 337654 Cherin etal. (2016) 13.8
Lutraeximia umbra Pantalla SBAU 337654 Cherin etal. (2016) 13.9
Lutra simplicidens Voigtstedt IQW 1966/7313 (Voi. 3525) Cherin (2017) 10.3
Lutra simplicidens Voigtstedt IQW 1966/7314 (Voi. 3532) Cherin (2017) 10.7
Lutra palaeindica Upper Siwalik NHM M 37151 Raghavan etal. (2007) 11.5
Lutra lutra Hoxne BM4936 Willemsen (1992) 12.3
Cyrnaonyx antiqua Tornewton Cave BM M 34370 Willemsen (1992) 11.2
Cyrnaonyx antiqua Tornewton Cave BM M 34373 Willemsen (1992) 11.3
Cyrnaonyx antiqua Tornewton Cave BM M 34374 Willemsen (1992) 11.6
Cyrnaonyx antiqua Tornewton Cave BM M 50707 Willemsen (1992) 11.0
Cyrnaonyx antiqua Verona M7 Willemsen (1992) 11.8
Fig. 5 Geographical distribution
of the body size estimates
Lutra lutra from Grotta Romanelli and the puzzling evolutionary history of Eurasian otter
1 3
diversification of ecological niches. The other fossil Euro-
pean otter, Lutraeximia umbra, according to Cherin etal.
(2016) had a strong preference for hard prey (molluscs), but
possessed intermediate features between fish specialist and
hard invertebrates dentition.
Finally, several scholars detected geographic-related
body size differences in extant otters, which have been
explained according to Bergmann’s ecogeographical rule
(Yom-Tov etal. 2006; Meiri etal. 2007), or advocating a
detrimental effect of environmental pressures and food avail-
ability (Yom-Tov etal. 2010). Despite the paucity of the
otter sample prevents definitive conclusions, body size esti-
mates of the fossil specimens would show no dimensional
clines across Europe, neither geographic, nor chronological
Homo‑otter interactions
The finding of the fossil otter in the deposit of Grotta
Romanelli opens another question that requires attention.
Although the studied specimen lacks both exact strati-
graphical and taphonomic information, in level G lithic
artifacts referred to Mousterian (Middle Paleolithic) were
also found, which document the human occupation of
the cave. The potential Homo-Lutra interactions during
Pleistocene are indeterminate, and in need of an in-depth
emphasis from both paleoecological and archaeozoologi-
cal points of view. Man exploited carnivorans for a wide
range of purposes, among which food and fur, but even for
bone artifact industry, symbolism, and ornamental pur-
poses (Mallye 2007, 2011). The Homo medium-to-small-
sized carnivoran interactions are difficult to investigate due
to the fragility of the skeletal bones (Kurtén 1968). Indeed,
the majority of the carnivoran records from archeologi-
cal deposits are represented by fragmentary craniodental,
metapodial and phalange remains. This is probably due
to selective human exploitation, since the skull and distal
portion of the limbs have lesser nutritional return. In addi-
tion, the carnivoran species are usually rare in archeologi-
cal deposits, as for instances Riparo Mochi (2.9%, Tagli-
acozzo etal. 2012), Arene Candide (27.7%, Cassoli and
Tagliacozzo 1994a), Riparo Fumane (16.8%, Cassoli and
Tagliacozzo 1994b), Grotta Maggiore di San Bernardino
(26.5%, Cassoli and Tagliacozzo 1994c) and upper com-
plex (level E–A) of Grotta Romanelli (26.9%, Tagliacozzo
2003). However, the presence of carnivorans and Homo
in the same deposit not necessarily indicates that these
taxa were hunted. Cassoli and Tagliacozzo (1994a), ana-
lyzing the fossil remains from the long sedimentary suc-
cession of Arene Candide, suggested that the majority of
the carnivoran fossil remains do not show cut mark and
often were found in anatomical connection. The authors
also identified bite marks in a few carnivoran specimens,
as well as some ungulate ones, suggesting therefore that
this cave was also seasonally frequented by large car-
nivorans (leopard or hyaena) or more stable occupied by
bear, where the prevalence of young individuals has been
interpreted as its possible den. Recently, a natural accu-
mulation of an abundant sample of L. lutra from Bliznets
Cave (Russian Far East) has been described, testifying that
otters can visit caves as well as perish in them(Alekseeva
andBaryshnikov2020). In this scenario, who hunted the
L. lutra from Grotta Romanelli cannot be ascertained, even
if the abundance of the Middle Paleolithic artifacts from
level G points to the Neanderthal as possible responsible,
which could have included otter in its diet.
The Eurasian otter from the late Middle Pleistocene of
Grotta Romanelli represents one of the earliest evidences
of the species, previously known only since the Holocene
onwards. The comparison analyses allow to exclude an attri-
bution to L. simplicidens and C. antiqua. L. simplicidens and
L. lutra have a similar size and share a fish-feeding type of
dentition, but they are distinct by the proportions of their
lower carnassial teeth and morphological features of the
upper carnassial. C. antiqua has a shellfish-feeding denti-
tion, broader and more robust than that of L. lutra. However,
here, we recognize that the two species overlap in chrono-
logical distribution, which urges to exercise caution in the
attribution of isolated remains.
The occurrence of the Eurasian otter from the “terre
rosse” of Grotta Romanelli provides valuable paleoeco-
logial information. L. lutra, nowadays, is mainly living in
fresh and clear running waters or lakes, and a certain degree
of adaptation to brackish water is possible. Most likely, a
hydrographic network could have been occurred in the area
in front of the cave, at present under the sea level.
Unfortunately, taphonomic data are not enough to recon-
struct who buried the mandible and how, at present the only
bone referable to the species, in the “terre rosse”. Anyway,
the occurrence of L. lutra at Grotta Romanelli suggests that
the taxon spreads into Europe earlier than supposed before,
at least at the beginning of the Late Pleistocene or during
the late Middle Pleistocene, replacing L. simplicidens and
coexisting with C. antiqua.
Genetic evidence resume L. lutra as closely related with
the extant L. sumatrana (Hung and Law 2016), and it has
been suggested that both derived from the Upper Siwalik L.
palaeindica (Willemsen 1992). However, despite the lack
of cranial material prevents conclusive comparisons, our
analyses reveal strong morphological affinities between L.
palaeindica and L. simplicidens.
B. Mecozzi etal.
1 3
Further discoveries will shed new light on systematics
and evolution of these carnivorans in the Quaternary of
Acknowledgments The authors are thankful for granting access to the
material, the kindness and availability of Francesca Alhaique, director
of Archaeozoology laboratory of the Museum of Civilizations (includes
the National Prehistoric Ethnographic Museum “Luigi Pigorini”). We
are deeply indebted to the Soprintendenza Archeologia, Belle Arti e
Paesaggio delle province di Brindisi, Lecce e Taranto for the permission
of the research and field activities (2015–2017 and 2018–2020, resp.
R. Sardella) (Maria Piccarreta, Laura Masiello and Serena Strafella).
Many people provided their competence and help during fieldwork: L.
Bellucci, A. Billi, M. Brilli, J. Conti, L. Forti, G. Gaglianone, B. Giac-
cio, F. Giustini, D.A. Iurino, F. Strani, D. Sigari and many students. N.
Ciccarese, T. De Santis, don P. Frisullo, A. Capraro and M. Rizzo for
their precious logistic support. Special thanks Luca Forti for the photo
of the Grotta Romanelli bay (Fig.1d). Finally, we thank the reviewers,
G.F. Baryshnikov and M. Sabol, who greatly improved this article with
constructive criticisms, comments, and suggestions. This work was
funded by the Grandi Scavi 2016 (Grant no: SA116154CD9592F3);
Grandi Scavi 2017 (Grant no: SA11715C81468801); Grandi
Scavi 2018 (Grant no: SA1181642D3B3C58); Grandi Scavi 2019
(Grant no: SA11916B513E7C4B); Grandi Scavi 2020 (Grant no:
Funding Open access funding provided by Università degli Studi di
Roma La Sapienza within the CRUI-CARE Agreement..
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... Legend: A-Aves; Am-Amphibia; Bt-Bone tools; C-Crustacea; E-Echinodermata; Ft-Flint tools; For-Foraminifera; H-Hearts; Hr-Human remains; Lt-Limestone tools; M-Mammal; mMal-marine Malacofauna; cMal-continental Malacofauna; Ost-Ostracods; P-Pisces; Pa-Portable Art; R-Reptilia. (References 23,24,34,35,[38][39][40][41][42] Thinly to medium layered sands, silts and clays with stone lines and lenses of matrix-supported angular to subangular fine to coarse-grained monogenic limestone gravels A,B,C,D,E Bt, Ft, H, Pa A, Am, E, Hr, M, cMal, mMal, P, R ...
... Due to the "colluvial" nature of the sediments inherited by the erosion of Interglacial leached soils ISU3 can be correlated to the early phases of climate deterioration that followed the Last Interglacial (i.e., MISs 5d-b). Fossils from ISU3 were recently debated and rhino remainss were attributed to Stephanorhinus hemitoechus 42 , canid specimens ascribed to Canis lupus 34 and otter material to Lutra lutra 39 . ISU3 and ISU4 are in turn buried by ISU5, the so-called "Terre Brune '' (due to their brownish colours) of Blanc 23 (Fig. 3, Fig. 4). ...
... The chronological attribution of ISU3 to the early Late Pleistocene supports their European extinction after MIS 5. The complete hemimandible of Lutra lutra (Linnaeus, 1758) found during Blanc's excavations 23,24 was recently described by Mecozzi et al. 39 confirming its taxonomic attribution. Since the European record of this carnivoran is quite scarce and mainly known from Holocene deposits 64 , its presence within ISU3 represents one of the earliest evidences of L. lutra in Europe. ...
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During the last century, Grotta Romanelli (Southern Italy) has been a reference site for the European Late Pleistocene stratigraphy, due to its geomorphological setting and archaeological and palaeontological content. The beginning of the sedimentation inside the cave was attributed to the Last Interglacial (MISs 5e) and the oldest unearthed evidence of human occupation, including remains of hearths, was therefore referred to the Middle Palaeolithic. Recent surveys and excavations produced new U/Th dates, palaeoenvironmental interpretation and a litho-, morpho- and chrono-stratigraphical reassessment, placing the oldest human frequentation of the cave between MIS 9 and MIS 7, therefore embracing Glacial and Interglacial cycles. These new data provide evidence that the sea reached the cave during the Middle Pleistocene and human occupation occurred long before MISs 5e and persisted beyond the Pleistocene- Holocene boundary.
... The archaeological and palaeontological ndings from Grotta Romanelli, hosted in museums and institutions across Italy, have been the subject of several studies, both con rming (26,27) or questioning (28, 29, 30) the chrono-stratigraphic setting proposed by Blanc (20). Recently, new excavations led to a reassessment of the stratigraphy of the uppermost part of the cave in lling (31) and a partial revision of some of the palaeontological remains (32,33,34,35). Furthermore, a critical review of the Last Interglacial PSLs' along the stable coasts of the Mediterranean Sea (11), indicated the highest Grotta Romanelli's notch as older than MISs 5e. ...
During the last century, Grotta Romanelli (Southern Italy) has been a reference site for the European Late Pleistocene stratigraphy, due to its geomorphological setting and archaeological and palaeontological content. The beginning of the sedimentation inside the cave was attributed to MISs 5e and the oldest unearthed evidence of human occupation, including remains of hearths, was therefore referred to the Middle Palaeolithic. Recent surveys and excavations produced new U/Th dates, palaeoenvironmental interpretation and a litho-, morpho- and chrono-stratigraphical reassessment, placing the oldest human frequentation of the cave between MIS 9 and MIS 7, therefore embracing Glacial and Interglacial cycles. These new data provide evidence that the sea reached the cave during the Middle Pleistocene and human occupation occurred long before MISs 5e and persisted beyond the Pleistocene- Holocene boundary.
... The spotted hyena becomes extensively represented since the late Middle Pleistocene. For instance, abundant samples were recovered from Bristie I (Lugli and Sala, 2000) and Melpignano (Vinuesa et al., 2016;Iannucci et al., 2020b;Mecozzi et al., 2021a), and the geographical range of the species reached the southeastern extremity of Italy at Grotta Romanelli (Sardella et al., 2018;Mecozzi et al., 2021b) and even Sicily, with the richest locality being San Teodoro (Mangano, 2011). The presumed evidence for Neandertal ritual behavior at Grotta Guattari was also reconsidered as the result of the activity of C. crocuta (Stiner, 1991). ...
The giant, short-faced hyena Pachycrocuta brevirostris was the largest Hyaenidae ever existed and the one that perfectly embodied the distinctive bone-cracking adaptations of this mammal family. Its dispersal into Europe is regarded as a biochronological marker of the Late Villafranchian at ~2.0 Ma, and its potential ecological interactions with other carnivorans and early Homo populations diffusing Out of Africa have given rise to extensive discussions. Nevertheless, our comprehension of the extinction of P. brevirostris remains vague. Here, we first critically evaluate the European fossil record of the species and then we review the whole Epivillafranchian and Galerian Hyaenidae record, including P. brevirostris, Crocuta crocuta and "Hyaena" prisca. Biometric comparisons with other extinct and extant bone-cracking hyenas are carried out. In contrast to a common view, we recognize that there is neither evidence of a persistence of P. brevirostris beyond the Early-Middle Pleistocene boundary, nor of a coexistence between the giant hyena and C. crocuta. The replacement between the two species, which was also accompanied by the arrival of "H." prisca, occurred at ~0.8 Ma and can serve as a marker of the Epivillafranchian–Galerian turnover, part of the faunal renewal that reflects the response of mammal communities to the Early–Middle Pleistocene Transition. Moreover, we clarified that Pliocrocuta perrieri and "H." prisca were different species, and that the latter was relatively more widespread than often assumed, being recorded from localities spanning in age almost the whole Middle Pleistocene and even the early Late Pleistocene.
... Within the terre brune layers, the palaeontological record is characterised by temperate species, mainly from steppe and grassland environments, along with North Atlantic and subarctic marine species, such as great auk (Pinguinus impennis) (see Sardella et al. 2018;Mecozzi et al. 2021). The lithic material totalled approximately 10 000 artefacts (with some 5800 pieces in Layer C), including small end scrapers, burins, finely retouched points on a blade, a bladelet or flake and many backed and truncated lithics (Sardella et al. 2019). ...
The Romanelli Cave in south-east Italy is an important reference point for the so-called ‘Mediterranean province’ of European Upper Palaeolithic art. Yet, the site has only recently been subject to a systematic investigation of its parietal and portable art. Starting in 2016, a project has recorded the cave's interior, discovering new parietal art. Here, the authors report on a selection of panels, featuring animal figures, geometric motifs and other marks, identifying the use of different types of tools and techniques, along with several activity phases. These panels are discussed with reference to radiocarbon dating of nearby deposits, posing questions about chronology, technology and wider connections between Upper Palaeolithic cave sites across western Eurasia.
... In addition to these taxa, the rare water buffalo Bubalus murrensis also occurs and spread into northern Europe during temperate climatic stages (Sardella, 2007 and references therein), whereas the Hemibos galerianus is quite scarce Martinez-Navarro & Palombo, 2007). Among cervids, in the early Middle Pleistocene European faunas Praemegaceros verticornis, Megaloceros savini and Dama roberti are recorded (e.g., Sardella et al., 2006;Sardella, 2007;Breda & Lis- project initiated during the 2015, whose preliminary results suggest that a possible late Middle Pleistocene age for the lower complex cannot be ruled out Mecozzi et al., 2021a). The narrownosed rhinoceros, S. hemitoechus is instead an important marker, as its earliest occurrence in Europe is dated approximately at 0.5 Ma (Caune de l'Arago, from levels chronologically referred to MIS 14; Moigne et al., 2006). ...
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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.
Conference Paper
Grotta Romanelli is a coastal cave inhabited by humans since the Middle Pleistocene and considered a symbol of the Palaeolithic period in Europe. The site, facing the Ionian Sea, is located in the administrative territory of the Castro (Lecce) municipality, at the south-eastern extremity of Apulia.This area, also known as Salentine Peninsula, documents the Quaternary human presence and climatic changes through rich archaeological, palaeontological and geological evidence. This richness favoured the establishment of several Regional Natural Parks to protect the natural landscape and its historical heritage, such as “Porto Selvaggio and Palude del Capitano”, “Isola di Sant’Andrea e Litorale di Punta Pizzo”, “Litorale di Ugento”, “Bosco and Paludi di Rauccio” and “Costa Otranto-Santa Maria di Leuca and Bosco di Tricase” where Grotta Romanelli is located.
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Procedeeding Book of the ProGEO SW Europe Regional Working Group. Virtual Conference on Geoconservation. March 30-31, 2022
Pollen analyses have been carried out on the infilling deposits of Grotta Romanelli (Apulia, Italy), a reference site for the Middle and Upper Palaeolithic of Italy. The analysis focused on Terre rosse, a fine unit till now ascribed to an interstadial phase following the Würm acme, and on the uppermost unit (Terre brune), recently dated to the latest Late Pleistocene-Early Holocene. Despite the diffuse barrenness and low pollen concentration of many levels, pollen data from Grotta Romanelli gives insights into the palaeoenvironmental setting of the deposits and their chronological attribution. The presence of Olea in all levels of Terre rosse strongly suggests their attribution to the Last Interglacial (Eemian), during which this plant was diffused in the Mediterranean area. The Terre brune deposition occurred when the environment was open, with rare trees and shrubs and prevailing steppe elements. This association reflects the climatic conditions of the Lateglacial, with evidence of both the warm interstadial Bølling/Allerød and the cold stadial Younger Dryas. Mediterranean, mesophilous and riparian arboreal elements are present, especially in the Early Holocene levels. Comparison with modern pollen material allowed some fossil grains, found in high amounts and in clusters, to be tentatively ascribed to the species Crithmum maritimum (Apiaceae), Muscari comosum and to Asparagus maritimus/Ornithogalum (Asparagaceae). The significant occurrence of such entomophilous plants reveals differential transportation inside the cave and, since most of them are edible and/or have medicinal properties, suggests an intentional introduction and possible use during time, by both Neanderthals and modern humans.
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The extant wild boar Sus scrofa has one of the largest geographical range of all mammals, and from its appearance in the late Early Pleistocene (Epivillafranchian) it is also widely represented in the European fossil record. Early forms of the species were larger than Late Pleistocene ones, but neither the chronology nor the causes of the size reduction have been thoroughly investigated. Here, we considered for the first time a large number of fossils from several late Middle Pleistocene to Early Holocene sites of the Apulian region (Italian Peninsula). In contrast to the supposed existence of a progressive trend towards small dimensions, morphometric comparisons and body mass estimates allow us to recognize several size oscillations during the late Middle Pleistocene-Early Holocene, with large forms occurring during interglacial stages and smaller ones during glacial stages. This suggests that fossil Apulian wild boar did not conform to Bergmann’s rule, that predicts larger size in colder climates due to the selective pressure towards lower surface area to volume ratio. Climate recrudescence may have played an indirect role in reducing the availability of trophic resources and hence promoting the observed pattern.
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Cranial remains of juvenile fossil rhinoceroses are rarely described in literature and very few is known about the ontogenetic development of their inner anatomy. In this study, we report the first CT based description of a juvenile braincase and its natural brain endocast of a late Middle Pleistocene Rhinocerotinae from Melpignano (Apulia, Italy). The specimen belongs to an individual about 12–18 months old, representing to date the youngest Pleistocene rhinoceros of Mediterranean Europe documented by neurocranial material. Through digital visualization methods the neurocranium has been restored and the anatomy of both the brain and the paranasal sinuses has been obtained and compared with those of juvenile and adult Pleistocene rhinoceroses. We evidence a different morphological development of the inner cranial anatomy in fossil and extant African species.
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In the last decades, many studies have focused on the description of fossil badger materials from Eurasia and several evolutionary hypotheses have been proposed. Nevertheless, the debate on taxonomy of the Late Villafranchian-Aurelian European badgers is still far from being solved and several species/subspecies were established over time. Herein, we described for the first time the craniodental and postcranial remains of Meles meles from Grotta Laceduzza (Apulia, Southern Italy), representing the largest sample of this taxon in the European Pleistocene record. Morphological and morphometric comparisons with fossils coming from the European Pleistocene sites were carried out; morphometric data were also compared with those of several extant populations of the European badger. The results of this work suggest that the badger fossil remains from the Mediterranean region can be considered as an ecomorphotype of this highly polymorphic species, showing a great morphological and morphometric variability throughout its wide geographical range. This variability is mainly expressed in some craniodental features and body size and could reflect local ecological adaptations, also linked to glacial/interglacial cycles.
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Fossil remains of felids from Geographical Society Cave and neighboring localities (Tigrovaya Cave, Malaya Pensau Cave, and Letuchiya Mysh Cave) in the Russian Far East are found to belong to four species: Panthera tigris, P. spelaea, P. pardus, and Lynx lynx. In Geographical Society Cave, the felid fossils are confined to deposits of the warm stage of the Late Pleistocene (MIS3). The simultaneous presence of Panthera tigris and P. spelaea seems to be unusual, the tiger remains being numerous whereas those of the cave lion are scant. There are differences between the Late Pleistocene tiger and the recent tiger in dental characters. P. tigris, most probably, migrated twice to Russian Far East from southern regions: in interstadial MIS3 and, subsequently, in the Holocene.
The paleontological collection from Geographical Society Cave located in southern part of Primorskii Territory is found to comprise 5 species of ursids and mustelids: Ursus arctos, Meles anakuma, Martes zibellina, Gulo gulo and Lutra lutra. Bone remains of brown bear (Ursus arctos) predominate; scant tooth-marks of large carnivores on their surfaces suggest bears to have been only occasional prey, dying mainly when overwintering in the cave. The presence of Asian badger (Meles anakuma) and true otter (Lutra lutra), whose findings are not known northwardly, provide the possibility to regard southern regions of the Russian Far East as a refuge, where these species survived during the Late Pleistocene.
The Romanelli Cave in south-east Italy is an important reference point for the so-called ‘Mediterranean province’ of European Upper Palaeolithic art. Yet, the site has only recently been subject to a systematic investigation of its parietal and portable art. Starting in 2016, a project has recorded the cave's interior, discovering new parietal art. Here, the authors report on a selection of panels, featuring animal figures, geometric motifs and other marks, identifying the use of different types of tools and techniques, along with several activity phases. These panels are discussed with reference to radiocarbon dating of nearby deposits, posing questions about chronology, technology and wider connections between Upper Palaeolithic cave sites across western Eurasia.
Caves as geosites structurally illustrate the strict dependence of human occupation on geological and geomorphological processes, playing a crucial role in the development of human civilisation. Grotta Romanelli embodies such a kind of geosite, being a coastal cave occupied by humans since the Middle Pleistocene and considered a symbol of the Palaeolithic period in Europe. Research on the cave, derived from the excavation activities carried out last century, consisted of a well-documented stratigraphic framework, abundant fossil remains and archaeological findings which included tools and rock art. The excavation activities stopped for about 40 years, hampering any new research on the cave. In 2015, new fieldwork was initiated and the multidisciplinary team immediately had to face several conservation issues linked to natural processes (erosion, degradation of the walls due to biodeteriogens) and human activities (mainly legal and illegal excavations). The use of 3D technologies to document the different phases of the research, from the field work to the digital reconstruction of fossil remains, has been extensively applied and represents an attempt to solve the issues of accessibility, education and sharing the heritage, which should be further implemented in the future.
In this paper, we present the results of the accelerator mass spectrometry radiocarbon (AMS ¹⁴ C) dating campaign performed on samples selected from different levels in Grotta Romanelli (Castro, Italy). Grotta Romanelli is one of the key sites for the chronology of Middle Pleistocene–Holocene in Mediterranean region. After the first excavation campaigns carried out in the first decades of the 1900s, the cave has been systematically re-excavated only since 2015. During the last excavation campaigns different faunal remains were selected and submitted for ¹⁴ C dating in order to confirm the chronology of the cave with a higher resolution. Isotopic ratio mass spectrometry (IRMS) measurements were also carried out on faunal remains.