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

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 & moyà-solà (1978). The supposed occurrence of Arvicola mosbachensis (scHmidtGen, 1911) pointed out by the latter authors, however, turned indeed to

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DOI: 10.1127/njgpa/2015/0460 0077-7749/2015/0460 $ 2.75

N. Jb. Geol. Paläont. Abh. 275/2 (2015), 151–161 ArticleStuttgart, February 2015E

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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)

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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.

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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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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, 1814Subfamily Erinaceinae FiscHer, 1814

Tribe Erinaceini FiscHer, 1814Genus Erinaceus linnaeus, 1758

Erinaceus cf. praeglacialis Brunner, 1933Fig. 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

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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, 1814Subfamily Crocidurinae milne-edwards, 1872

Genus Crocidura WaGler, 1832Crocidura 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. 432008 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 (Hermann, 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.

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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 - - 2.79CV-MC-620 4.16 - 3.44 2.92CV-MC-621 4.00 5.00 3.57 -CV-MC-622 - - 3.56 2.96CV-MC-623 4.56 5.05 - 3.07CV-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 2.69CV-MC-635 - - 3.47 3.38CV-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 - 3.11IPS-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)

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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).

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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 Galindo (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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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: [email protected] luis GiBert, Departament de Geoquímica, Petrologia i Prospecció Geològica, Facultat de Geologia, Universitat de Barcelona. c/ Martí Franques s/n, 08028 Barcelona, Spain;email: [email protected] Ferràndez, Departament d’Estratigrafia, Paleontologia i Geociències Marines, Facultat de Geologia, Universitat de Barcelona. c/ Martí Franques s/n, 08028 Barcelona, Spain;e-mail: [email protected] sevilla, Departamento de Paleontología, Facultad de Geología, Universidad Complutense de Madrid. c/ José Antonio Novais, 12, 28040 Madrid, Spain;e-mail: [email protected]

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Documents

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