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The insectivores (Soricidae, Erinaceidae; Eulipotyphla; Mammalia) from Cueva Victoria (Early Pleistocene, Murcia, Spain)

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Abstract and Figures

In the Early Pleistocene locality of Cueva Victoria, the insectivores are represented by several fossil teeth, mandibles and skull fragments. In the present work, we describe and provide measurements of these fossils for the first time. Insectivores show a rather low taxonomic diversity in Cueva Victoria, with only two different species. Considering the environmental requirements for their extant relatives, the presence of only one hedgehog (Erinaceus cf. praeglacialis Brunner, 1933), and one shrew (Crocidura kornfeldi Kormos, 1934) points to the existence of dry and warm conditions in the surrounding area of Cueva Victoria, unfavourable to other contemporary species of insectivores.
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The insectivores (Soricidae, Erinaceidae; Eulipotyphla; Mammalia)
from Cueva Victoria (Early Pleistocene, Murcia, Spain)
Marc Furió, Luis Gibert, Carles Ferràndez, and Paloma Sevilla
With 5 figures and 2 tables
Abstract: In the Early Pleistocene locality of Cueva Victoria, the insectivores are represented by
several fossil teeth, mandibles and skull fragments. In the present work, we describe and provide
measurements of these fossils for the first time. Insectivores show a rather low taxonomic diver-
sity in Cueva Victoria, with only two different species. Considering the environmental requirements
for their extant relatives, the presence of only one hedgehog (Erinaceus cf. praeglacialis Brunner,
1933), and one shrew (Crocidura kornfeldi Kormos, 1934) points to the existence of dry and warm
conditions in the surrounding area of Cueva Victoria, unfavourable to other contemporary species of
insectivores.
Key words: Crocidurinae, white-toothed shrew, hedgehog, Iberian Peninsula, glacial refuge, bioge-
ography, Quaternary.
1. Introduction
Cueva Victoria is a large karstic system with more than
three kilometers of galleries. It is a reference site of
the Early Pleistocene in Spain because it has provided
an impressive vertebrate assemblage, with thousands
of fossil elements of more than 70 different species of
vertebrates, some of them new to science (GiBert &
Ferràndez, research in progress). The most peculiar
element in this assemblage is the cercopithecid pri-
mate Theropithecus (Theropithecus) oswaldi leakeyi
Hopwood, 1934, which records the only unquestion-
able occurrence of this genus in Europe (Ferràndez et
al. 2014). The presence of Homo sp. in the assemblage
based on an intermediate phalanx has been discussed
(see GiBert et al. 2008; martínez-navarro et al. 2008,
and references therein), thus becoming controversial
for some authors. The fossil accumulation in the cave is
interpreted as a palaeoden of the hyena Pachycrocuta
brevirostris (aymard, 1846), which accounts for most
of the macromammal remains inside the cave (GiBert
et al. 1993; Ferràndez et al. 2011).
Within the small vertebrate assemblage from
Cueva Victoria, the contribution of the avifauna, with
almost forty different recorded species, is noteworthy
(sáncHez-marco 2004). The herpetofauna also helped
to extend the faunal list with at least eleven more spe-
cies of amphibians and reptiles, thus giving some in-
dications on the palaeoenvironmental conditions at the
moment of deposition (Blain et al. 2008). Regarding
small mammals, however, only some groups have
been studied in detail. The presence of rodents was al-
ready noticed in the first references to the site by pons-
moyà & mo-solà (1978). The supposed occurrence
of Arvicola mosbachensis (scHmidtGen, 1911) pointed
out by the latter authors, however, turned indeed to
©2015 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de
DOI: 10.1127/njgpa/2015/0460 0077-7749/2015/0460 $ 2.75
N. Jb. Geol. Paläont. Abh. 275/2 (2015), 151–161 Article
Stuttgart, February 2015
E
eschweizerbart_xxx
152 M. Furió et al.
be a new species of arvicoline named Allophaiomys
chalinei alcalde, aGustí & villalta, 1981. After its
original description, this species changed its generic
allocation several times, and it is nowadays consid-
ered the type species of the recently created genus
Victoriamys (martin 2012). Other than V. chalinei,
the rodent assemblage from Cueva Victoria was thor-
oughly described by aGustí (1982), in an attempt to
provide an age for the site, concluding that the faunal
association could be ascribed to ‘the Betfia phase in
the Biharian stage’ (i.e., close to 1.0 Ma). More recent-
ly, a revision of the material of lagomorphs from this
locality resulted in the description of a new species
of leporid, Oryctolagus giberti de marFà, 2008. All
these small vertebrate faunas are currently under revi-
sion, and they will be published elsewhere.
However, two important groups of small mammals,
bats and insectivores, were still to be systematically
studied. The former, the chiropterans, are currently
under revision by one of us (P. S.). According to gener-
al faunal lists, there were only two forms in the site of
the latter group, the insectivores, both identified at the
genus level: a shrew (Crocidura sp.) and a hedgehog
(Erinaceus sp.). Such a poorly diversified fauna was
surprising because fossil insectivores are frequently
more diverse in other similarly-aged micromammal
sites from Europe, rather suggesting a bias due to in-
sufficient sampling or studying of the insectivore as-
semblage. In the present work, we provide a detailed
description of these species, and the occurrences of
Erinaceus and Crocidura in Cueva Victoria are dis-
cussed from the biostratigraphical and palaeoenviron-
mental points of view.
2. Methods
The fossils from Cueva Victoria come from a fossilif-
erous breccia than infilled the karstic cavities during
the Early Pleistocene (Ferràndez et al. 1989; GiBert
et al. 2006). Most of the breccia was removed during
manganese-mining activities in the last century. Only
a small part of the breccia remained in situ, attached to
the walls and the ceiling and thus difficult to access. A
substantial part of the fossiliferous breccia excavated
by the miners still remains in blocks and loose soils in
the floor of the cavities. This removed sediment, more
accessible, is rich in fossil remains and has provided
a substantial part of the vertebrate fossils. Thus, the
collection of fossils from Cueva Victoria comes from
both in situ and re-excavated material. Anyway, strati-
graphic, sedimentologic, palaeontologic and tapho-
nomic studies have evidenced that the infill of the cave
was a continuous and unique event, and that all the
fossil remains, both in- and ex- situ localities have a
similar age (GiBert et al. in press). The breccia over-
lies autochthonous sediments (decalcification clay)
and is covered by a capping flowstone. This fossilif-
erous level has beed dated combining paleomagnetic
and 230Th/U methods at 0.99-0.78 Ma (GiBert et al. in
press).
The macrofossil elements were excavated mainly
in different locations in the main cavity, Sala Unión
(previously called Victoria I) and adjacent secondary
cavities (Fig. 1). A second room, Victoria II, about 100
m NNE away from the first one, was infilled from dif-
ferent original entrances in the Pleistocene and hence
it is less rich in fossils. The studied material was col-
lected in several prospections carried out in the late
1970s and earliest 1980s and from systematic excava-
tions performed from 1984 to 2011. The standard wet-
sieving techniques were only occasionally employed
due to logistic difficulties. Instead, dry-screening
process in situ was performed during excavations, us-
ing a sieve of 5 millimetres light-wide, that supplied
most of the fossil elements used in this study. Sieving
was applied to both in-situ deposits (Sala Victoria II,
Andamio Superior, Utrillas) and sediment removed by
the mining activities (Sala Unión, Descargador) (Fig.
1).
Acronyms and list of material studied of each species:
MAMC – Museo Arqueológico Municipal de Cartagena;
MGB – Museu de Geologia de Barcelona; ICP – Insitut
Català de Paleontologia Miquel Crusafont.
Crocidura kornfeldi
MAMC – Utrillas: CV-MC-600 (Hemimandible (R) m1-
m3); CV-MC-601 (Hemimandible (R) a1-m2); CV-MC-602
(Hemimandible (R) a1-m3); CV-MC-603 (Hemimandible
(L) p4-m2); CV-MC-604 (Hemimaxilar (R) P4-M2); Sala
Unión: CV-MC-606 (Maxillar with I1, A3 and P4 (L));
CV-MC-607 (Maxillar with A3-M2 (L) + A1-M1 (R)); CV-
MC-608 (Maxillar with A1-M1 and M3 (L) + A3 and M1-
M3 (R)); CV-MC-609 (Maxillar with A1-M3 (L) + I1-A1
(R)); CV-MC-610 (Hemimaxillar (L) A1-M1); CV-MC-611
(Hemimaxillar (L) A3-M2); CV-MC-612 (Hemimaxillar
(R) P4-M2); CV-MC-613 (Hemimaxillar (R) P4-M3);
CV-MC-614 (Hemimaxillar (R) A2-P4); CV-MC-615
(Hemimaxillar (R) P4-M1); CV-MC-616 (Hemimandible
(L) m1); CV-MC-617 (Hemimandible (L) p4 m3); CV-
MC-618 (Hemimandible (R) m1-m3); CV-MC-619
(Hemimandible (R) i1-m3); CV-MC-620 (Hemimandible
(R) i1-m3); CV-MC-621 (Hemimandible (L) m1-m3);
CV-MC-622 (Hemimandible (L) i1-m3); CV-MC-623
(Hemimandible (L) i1-m2); CV-MC-624 (Hemimandible (R)
eschweizerbart_xxx
The insectivores (Soricidae, Erinaceidae; Eulipotyphla; Mammalia) from Cueva Victoria 153
m1); CV-MC-625 (Hemimandible (R) p4-m3); CV-MC-626
(Hemimandible (R) p4-m3); CV-MC-627 (Hemimandible
(L) p4-m1); CV-MC-628 (Hemimandible (R) m1-m3);
CV-MC-629 (Hemimandible (L) m1-m3); CV-MC-630
(Hemimandible (L) m1); CV-MC-631 (Hemimandible (L)
p4-m3); CV-MC-632 (Hemimandible (R) m1-m3); CV-
MC-633 (Hemimandible (L)); CV-MC-634 (Hemimandible
(L) i1-m3); CV-MC-635 (Hemimandible (R) i1 + p4-m3);
CV-MC-636 (Hemimandible (L) p4-m2); CV-MC-637
(Hemimandible (R) m2); CV-MC-638 (Hemimandible
(R) p4-m3); CV-MC-639 (Hemimandible (R) m1-
m3); CV-MC-640 (Humerus); CV-MC-641 (Humerus);
CV-MC-642 (Humerus); CV-MC-643 (Humerus);
CV-MC-644 (Humerus); CV-MC-645 (Humerus);
Fig. 1. Plan view of the cave system with the outcrops where the fossils come from. Sala Unión includes both types of ma-
terial, ‘in situ’ and ‘ex situ’. The right-bottom part of the figure indicates the geographic situation of Cueva Victoria in the
Iberian Peninsula.
eschweizerbart_xxx
154 M. Furió et al.
Fig. 2. Measurements taken in the dental elements of Erinaceus described in this paper. The reference line in P4 joins the
points of maximum convexity of the anterolabial and posterolabial zones. The reference line in M1 and M2 joins the points
of maximum convexity of the anterolabial and the anterolingual zones. The reference line in m3 is tangent to the most
lingual point, and parallel to the longitudinal axis of the tooth. The parameters measured in each element are parallel or
perpendicular to their corresponding reference lines. Taken from Furió (2007a).
Fig. 3. Measurements taken on the hemimandibles of Crocidura kornfeldi from Cueva Victoria.
Fig. 4. Left hemimandible of Erinaceus cf. praeglacialis from Cueva Victoria (left; CV-MC-663, MAMC) compared with
the same element in the Recent species E. europaeus (right; ICP). Scale bar: 0.5 cm.
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The insectivores (Soricidae, Erinaceidae; Eulipotyphla; Mammalia) from Cueva Victoria 155
CV-MC-646 (Humerus); CV-MC-647 (Humerus);
CV-MC-648 (Humerus); CV-MC-649 (Humerus); CV-
MC-650 (Humerus); CV-MC-651 (Humerus); CV-
MC-652 (Humerus); CV-MC-653 (Humerus); CV-MC-654
(Pelvis); CV-MC-655 (Femur); CV-MC-656 (Tibiofibula);
Sala Victoria II: CV-MC-605 (Humerus); CV-MC-667
(Hemimandible (L) m1-m2); CV-MC-668 (Hemimandible
(R) m1); CV-MC-669 (Hemimandible (L)); CV-MC-670
(Hemimandible (L) p4-m2); CV-MC-671 (Hemimandible
(R) m1-m3); CV-MC-672 (Hemimandible (L) p4-m3)
ICP – Sala Unión: IPS-20998 (Maxillar with A2-M3 (R)
and A2+P4 (L)); IPS-21000 (Hemimandible (L) i1-m1); IPS-
46863 (Hemimandible (L)); IPS-46864 (Hemimandible (L)
m1-m3); IPS-46865 (Hemimandible (L) m1); IPS-46866
(Hemimandible (R) a1-m3); IPS-46867 (Hemimandible
(L) m1 m3); IPS-46868 (Hemimandible (R)); IPS-46869
(Hemimandible (R) p4-m2); IPS-46870 (m1 (L)); IPS 46871
(m1 (L)); IPS-46872 (m3 (L)); IPS-46873 (m2 (L)); IPS-
46874 (Maxillar with I1-A1 and P4-M1
(R) and I1-A1 (L)); IPS-46875 (I1 (R)); IPS-46876 (a1 (R));
IPS-46877 (A1 (R))
MGB – MGB-6157 (Hemimandible (R))
Erinaceus cf. praeglacialis
MAMC – Descargador: CV-MC-664 (Mandibular frag-
ment (R)); Sala Unión Este: CV-MC-661 (Mandibular
fragment (L)); CV-MC-662 (Mandibular fragment (R));
Sala Unión Oeste: CV-MC-660 (Mandibular fragment
(R)); Sala Unión: CV-MC-658 (Hemimaxillar (L) P4-M2);
CV-MC-659 (Mandibular fragment (L)); CV-MC-663
(Hemimandible (L)); CV-MC-665 (Mandibular fragment);
CV-MC-666 (Mandibular fragment (L) Juvenile with i1
and p4); Andamio Superior E: CV-MC-673 (M1 (L)); CV-
MC-674 (m3 (L))
Nomenclature and measurements of the erinaceids follow
Furió (2007a). The way of measuring P4, M1 and m3 is de-
tailed in Fig. 2. The nomenclature and the measurements of
Crocidura follow reumer (1984), but only some measure-
ments of the mandible (Length, Height, Length m1-m3, and
Length of the Lower Incisor) have been considered signifi-
cant, as detailed in Fig. 3.
Table 1. Dental measurements (in mm.) of Erinaceus cf.
praeglacialis from Cueva Victoria.
Specimen Element Length Width
CV-MC-658 P4 4.98 5.23
CV-MC-673 M1 5.61 5.84
CV-MC-658 M1 6.70 6.53
CV-MC-658 M2 4.89 6.58
CV-MC-674 m3 2.52 2.35
3. Systematic palaeontology
Family Erinaceidae FiscHer, 1814
Subfamily Erinaceinae FiscHer, 1814
Tribe Erinaceini FiscHer, 1814
Genus Erinaceus linnaeus, 1758
Erinaceus cf. praeglacialis Brunner, 1933
Fig. 4, Table 1
1981 Erinaceus sp. – carBonell et al., p. 49.
2006 Erinaceus cf. europaeus. – GiBert et al., p. 43.
2008 Erinaceus cf. E. europaeus. – Blain et al., p. 347.
Remarks: The material of Erinaceus is scanty in Cueva
Victoria. No more teeth than those of the specimen CV-
MC-658 (a P4, a M1 and a M2), CV-MC-673 (a M1), CV-
MC-674 (a m3), and the not yet erupted i1 and p4 of the
specimen CV-MC-666 have been found (measurements of
these elements are given in Table 1). This is really a scarce
sample, hampering precise identification of the species.
However, the ascription to the genus Atelerix (a possible
option considering the presence of other African genera
such as Theropithecus or Crocidura) is confidently re-
jected because in the specimen CV-MC-658 there is only
one alveolus for the P2, a characteristic trait of the genus
Erinaceus (corBet 1988: table 3). Moreover, the specimen
CV-MC-673 (M1) has a long posterior cingulum ending at
its posterolingual corner, unlike Atelerix, in which it usually
ends at the level of the hypocone tip (Butler & Greenwood
1973: table 6).
The tentative ascription to E. praeglacialis is based on
the data provided by these few elements and the morpholo-
gy of the left hemimandible CV-MC-663. The most evident
trait observed is that the erinaceid present in Cueva Victoria
is somewhat larger than the recent species E. europaeus
Linnaeus 1758 (18% in height of the coronoid process; > 8%
mandibular length; see Figure 4). Other than size, the fossil
species differs from the recent one by its longer metacrista
in M1 with a less labial and more posterior orientation in the
fossil than in the recent form. This character was pointed
out in the original description of the species (Brunner 1933)
and also considered by rzeBiK-KowalsKa (2000) as diag-
nostic of E. praeglacialis.
Some other characters never referred in literature are
worth mentioning. For instance, the ventral border of the
horizontal ramus is less curved in the material from Cueva
Victoria than in E. europaeus (Fig. 4). There is also a no-
ticeable difference in the curvature of the posterior margin,
between the uppermost side of the coronoid process and the
articular condyle. Moreover, the ridge delimiting the ante-
rior border of the temporal fossa is stouter in the fossil form
than in the extant species. It is also noteworthy that the an-
terior margin of the coronoid process is somewhat straighter
in Erinaceus from Cueva Victoria than in the recent mate-
rial.
It must be considered that the taxonomy of the fossil
forms of Erinaceus is in urgent need of revision, as there
are no unified criteria to differentiate the Plio-Pleistocene
eschweizerbart_xxx
156 M. Furió et al.
forms from Europe. Therefore, the ascription of the ma-
terial from Cueva Victoria to the species E. praeglacialis
must be considered tentative, and it is pending a review of
Plio-Pleistocene Erinaceus that the first author (M.F.) is
currently carrying out.
Family Soricidae FiscHer, 1814
Subfamily Crocidurinae milne-edwards, 1872
Genus Crocidura WaGler, 1832
Crocidura kornfeldi Kormos, 1934
Fig. 5, Table 2
1978 Crocidura kornfeldi Kormos. pons-moyà &
moyà-solà, p. 54.
1981 Crocidura sp. – carBonell et al., p. 49.
2006 Crocidura kornfeldi. – GiBert et al., p. 43
2008 Crocidura kornfeldi. – Blain et al., p. 347.
Remarks: Good descriptions of the dental and mandibular
characters of C. kornfeldi can be found in reumer (1984),
and roFes & cuenca Bescós (2011), among others. Thus,
descriptions are here limited to highlight the most distinc-
tive characters regarding or differentiating such material. In
general terms, all the dental, mandibular and maxillar ele-
ments from Cueva Victoria are smaller than those of recent
C. russula (Hermann, 1780) and C. suaveolens (pallas,
1811). Comparing the mandibular height (H) of C. kornfeldi
from CV with the measurements of recent species provided
by saint-Girons et al. (1979), only the range of C. leuco-
don (Her mann, 1780) falls within that of the fossil species.
Morphologically, the most distinctive trait differentiating
the fossil species from the recent European ones is the posi-
tion of the protocone in P4, which is not immediately at the
anterolingual corner, but somewhat more labially displaced.
A significant difference with previous records of C. ko-
rnfeldi is found in the relative size of the upper antemolars.
In the material from CV, the A1 is much larger than A2, and
A3 is only slightly smaller (or equal in size) than A2. This is
somewhat different than the description by reumer (1984)
for the Hungarian material, where the A2 is a bit smaller
than the A3.
When compared with the measurements in reumer
(1984), the material from CV closely fits with C. kornfeldi
(Table 2) from Villány 3, but is considerably smaller than
the material from Osztramos 3/2. Our measurements are
also in good agreement with the ranges provided by roFes
& cuenca-Bescós (2011) for the mandibles of C. kornfeldi
from Sima del Elefante-Atapuerca.
4. Discussion
4.1. Representativity of the assemblage
The sampling technique employed has obviously re-
sulted in a bias of the fossil elements recovered in
Fig. 5. Crocidura kornfeldi, maxilla (a; CV-MC-607, MAMC) and mandible (b; CV-MC-635, MAMC) compared with the
same elements in its recent relative C. russula (c, d; ICP). Notice the different size of both species. Scale bar: 0.5 cm.
eschweizerbart_xxx
The insectivores (Soricidae, Erinaceidae; Eulipotyphla; Mammalia) from Cueva Victoria 157
Cueva Victoria. It is evident that a sieve with a light
of 5 mm could only retain the mandibles and partial
skulls of these insectivores. Not many loose teeth have
been found, except for the two teeth of Erinaceus. The
few isolated teeth of Crocidura from the ICP collec-
tions more likely belong to rather complete remains
disarticulated after their storage.
Nevertheless, the observed taxonomic diversity of
insectivores at this site does not seem to be affected by
the sampling methods. If that were the case, other simi-
Table 2. Mandibular measurements (in mm.) of Crocidura kornfeldi from Cueva Victoria. M: mean; R: range; SD: standard
deviation; N: number of specimens.
Specimen Mand.
Height
Mand.
Length
Length
m1-m3
Length
i1
CV-MC-600 4.06 4.91 3.42 -
CV-MC-601 4.20 - - -
CV-MC-602 - - 3.53 -
CV-MC-603 4.29 - - -
CV-MC-616 4.00 5.18 - -
CV-MC-617 4.45 5.12 3.67 -
CV-MC-618 4.43 5.12 3.57 -
CV-MC-619 4.02 - - 2.79
CV-MC-620 4.16 - 3.44 2.92
CV-MC-621 4.00 5.00 3.57 -
CV-MC-622 - - 3.56 2.96
CV-MC-623 4.56 5.05 - 3.07
CV-MC-624 4.16 5.48 - -
CV-MC-625 4.23 5.06 3.44 -
CV-MC-626 4.08 5.10 3.50 -
CV-MC-628 4.58 5.52 3.59 -
CV-MC-629 4.35 5.11 3.44 -
CV-MC-630 4.47 4.96 - -
CV-MC-631 4.14 4.97 3.39 -
CV-MC-632 4.37 4.99 3.47 -
CV-MC-634 4.20 5.00 3.34 2.69
CV-MC-635 - - 3.47 3.38
CV-MC-637 4.22 4.80 - -
CV-MC-638 4.08 4.83 3.52 -
CV-MC-639 4.32 5.15 3.49 -
CV-MC-667 4.49 5.30 - -
CV-MC-668 4.12 5.28 - -
CV-MC-669 - 5.41 - -
CV-MC-670 - 5.24 - -
CV-MC-671 4.23 5.06 3.52 -
CV-MC-672 4.18 5.28 3.38 -
IPS-21000 4.29 5.38 - 3.11
IPS-46863 4.05 - - -
IPS-46864 3.91 5.12 3.67 -
IPS-46865 4.32 5.23 - -
IPS-46866 4.36 5.26 3.66 -
IPS-46867 3.95 4.87 3.58 -
IPS-46868 3.97 - - -
IPS-46869 4.01 5.00 3.40 -
MGB-6157 4.10 5.23 - -
M (R)
SD (N)
4.20 (3.91-4.58)
0.180 (35)
5.12 (4.91-5.52)
0.182 (31)
3.50 (3.34-3.67)
0.094 (23)
2.98 (2.69-3.38)
0.226 (7)
eschweizerbart_xxx
158 M. Furió et al.
lar-sized elements of different taxa would be expected.
Moreover, Cueva Victoria includes several outcrops
(Utrillas, Sala Unión, Sala Victoria II, Descargador
and Andamio Superior) of the fossiliferous breccia in
different parts of the cave, all of which show the same
limited diversity (Fig. 1). The possibility that other
species lived in the surrounding areas in the past but
they were not recorded in the fossil sample is quite un-
likely. Thus, the large sample available, obtained from
different points, must be taken as a good estimation of
the real paleodiversity of insectivores during the mo-
ment of the karstic infilling. A preferential distribution
of Crocidura kornfeldi or Erinaceus cf. praeglacialis
is not either discerned, and both are quite uniformly
represented in the breccias where they come from.
4.2. Biostratigraphy
The presence of Crocidura in Europe is documented
from the Pliocene onwards (reumer 1984). In Spain,
Crocidura had been apparently reported in several
Pliocene sites (van den HoeK ostende & Furió 2005).
However, a revision of the material revealed that the
white-toothed shrew present in these Pliocene lo-
calities was indeed Myosorex meini (Furió et al.
2007), so the oldest real occurrences of Crocidura
in the Spanish fossil record correspond to the Early
Pleistocene (Furió 2007b). The arrival of the genus to
the Iberian Peninsula can be placed between 1.8 and
1.3 Ma. (Furió 2007b).
The specific ascription of the oldest representatives
of the genus in Spain has been uncertain for a long time.
However, roFes & cuenca-Bescós (2011) determined
that the form present in the Early Pleistocene site of
Sima del Elefante, one of the oldest Spanish sites with
Crocidura, was in fact C. kornfeldi. The morphologi-
cal and biometrical analyses indicate that the material
from Cueva Victoria belongs to this species as well.
Considering these two occurrences, C. kornfeldi was
likely the first representative of Crocidura to colonize
the Iberian Peninsula.
The fossil record of Erinaceus in the Iberian
Peninsula is rather scarce. Its occurrences seem ran-
domly distributed in time and space. In the compiled
data by van den HoeK ostende & Furió (2005), the
oldest record of the genus corresponds to Aljezar B,
an MN 12 (Turolian) locality from Teruel. However,
this taxonomic ascription must be considered prelimi-
nary, as it was explicitly noticed that further research
on the material from this locality is pending (van den
HoeK ostende & Furió 2005). The next younger oc-
currences of Erinaceus in Spain come from the Early
Pliocene locality of La Gloria 4 (MN 14, Teruel) and
the Late Pliocene sites of Layna and Sarrión (MN 15)
and Escorihuela (MN 16). In the Early Pleistocene
sites, like Fuente Nueva 3, Barranco León, Sima
del Elefante and Gran Dolina (Early Pleistocene),
the genus is always represented by sparse fossil ele-
ments. In Fonelas P-1 (MN 17 - Early Pleistocene),
laplana & Blain (2008) documented the presence of
an Erinaceinae. The fossils from this locality were at-
tributed to a smaller and more slender species than the
recent E. europaeus, but similar to the erinaceid from
La Puebla de Valverde identified as Postpalerinaceus
cf. vireti by crocHet & Heinz (1971). If the observa-
tion of laplana & Blain (2008) is correct, the ascrip-
tion to the genus Postpalerinaceus is quite unlikely,
because Postpalerinaceus species are usually larger
than those of Erinaceus. Moreover, Postpalerinaceus
seems to be a genus exclusively restricted to Miocene
sites. Therefore, despite not yet determined at the ge-
nus level, the remains from Fonelas P-1 might be better
ascribed to Erinaceus as well. Yet the specific identi-
fication deserves further research, these observations
seem sufficient to identify this species from Fonelas
P-1 as different to the one present in Cueva Victoria.
Hitherto, the only similar form to that of Cueva
Victoria is E. cf. praeglacialis from the Early
Pleistocene localities of Fuente Nueva 3 and Barranco
León (Orce, Granada) described in Furió (2007a). This
limited range of E. praeglacialis (or similar forms) in
Spain agrees quite well with its occurrences in the rest
of Europe, where it is found in Early Pleistocene and
early Middle Pleistocene sites (Furió 2007a).
4.3. Paleoecology
The genus Crocidura is frequently regarded as in-
dicative of warm environments (reumer 1984, 1995;
rzeBiK-KowalsKa 1995; Furió 2007a; roFes &
cuenca-Bescós 2011). Although the genus is widely
distributed and adapted to all kind of environments
in Africa, its European counterparts mostly prefer
dry and rocky zones. In fact, the northern boundary
of its geographical distribution oscillated latitudinally
several times during the Early Pleistocene, probably
influenced by the alternation of cooling and warming
periods (reumer 1984; rzeBiK-KowalsKa 1995).
Erinaceus is a genus with a clear preference for mild
climates. Nowadays, the European forms avoid high lat-
itudes (i.e., above 53 °N), their distribution coinciding
with the extension of deciduous forests (corBet 1988).
eschweizerbart_xxx
The insectivores (Soricidae, Erinaceidae; Eulipotyphla; Mammalia) from Cueva Victoria 159
Some molecular studies indicate that the European
populations of Erinaceus could have been subjected to
cyclical restrictions to glacial refugia and interglacial
expansion during the Pleistocene (seddon et al. 2001;
sommer 2007), as happened with many other animals
and plants. During the glacial periods, the Iberian
Peninsula acted as a refuge for some of these spe-
cies adapted to rather mild climates, in a similar way
to the latitudinal shift showed by the fossil record of
Crocidura (santucci et al. 1998; sommer 2007).
The record of these two genera in Cueva Victoria
reinforces the role of the Iberian Peninsula as glacial
refuge for them during the Pleistocene. Unfortunately,
the southern location of Cueva Victoria does not permit
a refinement of the refuge area. Whereas the northern
and central areas of the Iberian Peninsula might have
been unfavourable for the two genera during some cold
periods, the more meridional latitude of Cueva Victoria
could have provided a milder climate for the survival
of both, independently from the global trend.
5. Conclusions
The systematic revision of the insectivore material
from Cueva Victoria stored in MAMC, MGB and
ICP, provides sound evidence that only two species
of insectivores are present in the fossil assemblage of
Cueva Victoria: the shrew Crocidura kornfeldi, and
the hedgehog Erinaceus cf. praeglacialis. This is a
quite short list of insectivores for an Early Pleistocene
karstic site that has yielded abundant material from
other small mammal groups. Apparently, there is no
reason to assume a bias in the taxonomic representa-
tion due to the sampling methods employed. These two
species are characteristic of the Early Pleistocene of
the Iberian Peninsula, yet their finds extend somewhat
longer in time in other European countries. Both gen-
era, Erinaceus and Crocidura, are frequently referred
as palaeoenvironmental indicators of rather warm or
dry conditions. This interpretation fits well with the
poor diversity of the site, which suggests rather in-
hospitable conditions, not dissimilar to the habitat of
Theropithecus today.
Acknowledgements
This work has been supported by the Ministerio de Economía
y Competitividad (GCL2011-28681) and the Generalitat
de Catalunya (2009 SGR 754 GRC, 2014 SGR 416 GRC,
and 2014 SGR 251). maría comas (MAMC), laura celià
and Jordi Galin do (ICP) and Julio Gómez-alBa (MGB)
are greatly acknowledged for permitting the authors to
check the collections. Special thanks go to david m. alBa
(ICP) for helping to take the photographs of Erinaceus and
Crocidura, to Jérôme prieto (Univ. Munich) for providing
essential literature and translating some German texts, and
to Consorcio Sierra Minera, Ayuntamiento de Cartagena,
Servicio de Patrimonio de la Comunidad de Murcia and
Museo Arqueológico Municipal de Cartagena, who fa-
cilitated the excavations and the study of the material. The
comments of Drs. emmanuel desclaux, BarBara rzeBiK-
KowalsKa and lars w. van den HoeK ostende helped to
improve the original manuscript. Dr. Günter scHweiGert is
acknowledged for his efficient and rapid management of the
present work as Editor of NJGP.
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eschweizerbart_xxx
The insectivores (Soricidae, Erinaceidae; Eulipotyphla; Mammalia) from Cueva Victoria 161
Manuscript received: September 19th, 1014.
Revised version accepted by the Stuttgart editor: October
23rd, 2014.
Addresses of the authors:
marc Furió (corresponding author), Institut Català de
Paleontologia Miquel Crusafont, Edifici Z (ICTA-ICP),
c/ de les Columnes, Campus de la UAB, Cerdanyola del
Vallès, 08193 Barcelona, Spain;
e-mail: marc.furio@icp.cat
luis GiBert, Departament de Geoquímica, Petrologia i
Prospecció Geològica, Facultat de Geologia, Universitat de
Barcelona. c/ Martí Franques s/n, 08028 Barcelona, Spain;
email: lgibert@ub.edu
carles Ferràndez, Departament d’Estratigrafia,
Paleontologia i Geociències Marines, Facultat de Geologia,
Universitat de Barcelona. c/ Martí Franques s/n, 08028
Barcelona, Spain;
e-mail: carlesferrandez@ub.edu
paloma sevilla, Departamento de Paleontología, Facultad
de Geología, Universidad Complutense de Madrid. c/ José
Antonio Novais, 12, 28040 Madrid, Spain;
e-mail: psevilla@ucm.es
eschweizerbart_xxx
eschweizerbart_xxx
... or Erinaceus sp. (Adrover, 1974;Mein et al., 1990;Van den Hoek Ostende and Furió , 2005;Furió et al., 2015Furió et al., , 2018Piñ ero et al., 2017), although the latter genus assignment should be regarded as uncertain. The youngest known record of pre-modern (i.e., 'non-Erinaceus') erinaceines is an occurrence of Postpalerinaceus (P. ...
... cf. vireti), as described for La Puebla de Valverde (MN17, Sarrió n depression); 1.5 Ma: entry of undisputed Erinaceus in the Lower Pleistocene (with various records across Spain; Furió et al., 2015Furió et al., , 2018. ...
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