Early reliable evidence of the Etruscan shrew (Suncus etruscus) in southwestern Europe...

Quaternary Science Reviews 250 (2020) 106690

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Early reliable evidence of the Etruscan shrew (Suncus etruscus) in southwestern Europe during ancient times. Reconstructing its dispersal process along the Mediterranean Basin

sar Laplana b, Paloma Sevilla a Angel C. Domínguez García a, *, Ce mica, Estratigrafía y Paleontología, Facultad de Ciencias Geolo gicas, Universidad Complutense de Madrid, Jos Departamento de Geodina e Antonio Novais, 12, 28040, Madrid, Spain b de Henares, Spain gico Regional de la Comunidad de Madrid, Plaza de las Bernardas, s/n, 28801, Alcala Museo Arqueolo a

a r t i c l e i n f o

a b s t r a c t

Article history: Received 29 April 2020 Received in revised form 6 October 2020 Accepted 28 October 2020 Available online xxx

This paper reports a new and well dated record of the Etruscan shrew (Suncus etruscus), which is the smallest extant terrestrial mammal, and provides a comprehensive review of the fossil record of this species. Three remains were found in the small vertebrate assemblage of the Estrecho Cave (Cuenca, Spain), which dates from 2310e2290 and 2272e2149 years cal BP. The morphometric analysis performed provided unequivocal traits of this species. We analyse this new record of the Etruscan shrew together with a review of its fossil records along the Mediterranean Basin. Because of the scarcity of reliable records of this species, its origin and palaeobiogeographical history remain still unclear. According to available data, the Etruscan shrew populations that colonised Europe were most probably originated in Asia, where it was present at least since the Middle Pleistocene. Its dispersal process across the Mediterranean Basin shows a pattern in which it progressively extended its distribution towards the west from mid-4th millennium BP onwards. This process most certainly involved the accidental translocation of the species by humans across the Mediterranean through navigation routes, resulting in its introduction into most of the large Mediterranean islands, besides the mainland of Europe and North Africa. © 2020 Elsevier Ltd. All rights reserved.

Keywords: Pigmy white-toothed shrew Holocene Estrecho cave Colonisation Ancient navigation

Credit author statement

Angel C. Domínguez García: Conceptualization, Methodology, Formal analysis, Investigation, Writing - original draft, Visualiza sar Laplana: Conceptualization, Validation, Formal analysis, tion. Ce Investigation, Writing- Review & Editing, Supervision. Paloma Sevilla: Conceptualization, Validation, Writing- Review & Editing, Supervision.

1. Introduction The pigmy white-toothed shrew or Etruscan shrew, Suncus etruscus (Savi, 1822) (Eulipotyphla, Soricidae), is one of the smallest pez-Fuster, 2007; living mammals with 1.2e2.7 g of weight (Lo Burgin and He, 2018). It has one of the largest distribution ranges

* Corresponding author.

E-mail addresses: angelcdo@ucm.es (A.C. Domínguez García), cesar.laplana@ gmail.com (C. Laplana), psevilla@ucm.es (P. Sevilla). https://doi.org/10.1016/j.quascirev.2020.106690 0277-3791/© 2020 Elsevier Ltd. All rights reserved.

among “insectivore” mammals, being widespread but scattered from Southern Europe and North Africa to Southeast Asia. It is also present in Madagascar and in the island of Tenerife. In Europe, it is restrained to the Mediterranean climate zone including most pezMediterranean islands (Blanco, 1998; Spitzenberger, 1990; Lo Fuster, 2007; Aulagnier et al., 2017; Burgin and He, 2018).

The genus Suncus belongs to the subfamily Crocidurinae and comprises 19 species inhabiting Eurasia and Africa (Burgin and He, 2018). It is mainly an Indomalayan genus, and therefore probably originated in southern Asia, where it subsequently diversified and spread into Africa and Europe (Heim de Balsac and Lamotte, 1957; Butler et al., 1989; Butler, 1998; Jenkins et al., 1998; Burgin and He, 2018), although some studies suggest an African origin for the rouil et al., 2001). The genus (Meester, 1953; McLellan, 1994; Que fossil record seemingly supports an African origin, due to the higher number of species in this continent and the abundance of fossil occurrences (Butler, 1998). However, a recent study describes the oldest known member of the genus in the Siwalik deposits of the Potwar Plateu, Pakistan, at 10.5 Ma (Flynn et al., 2020).

Multiple efforts to clarify the systematics and phylogeography of

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it is commonly accepted that the species colonised Europe during the Holocene, favoured by human activities (Rzebik-Kowalska, rouil et al., 2001; Vigne, 1999, 2014; 1998; Dobson, 1998; Que et al., 2018; Vigne and Pascal., 2003; Garrido-García, 2008; Furio Domínguez García et al., 2019a). However, both the precise age and the details of how this process took place remain still unclear.

Several authors claim that S. etruscus was introduced in Europe rouil et al., 2001; from North Africa (Rzebik-Kowalska, 1998; Que et al., 2018), whereas others propose an Garrido-García, 2008; Furio alternative scenario in which this species may have been introduced both in Europe and in North Africa from Eastern Mediterranean (Dobson, 1998; Domínguez García et al., 2019a). In any case, during the Holocene, several human-mediated biological invasions or translocations (intentional or accidental introduction of organisms into new ecosystems by humans) across the Mediterranean Basin have occurred related to the development of navigation techniques (Audoin-Rozeau and Vigne, 1994; Dobson, 1998; Vigne, 1999; Cucchi et al., 2005; Garrido-García, 2008; Domínguez García et al., 2019a).

In this study we report a Holocene record of S. etruscus found in the centre of the Iberian Peninsula, that sheds light on the palaeogeographical history of the species after putting together this record with the available records from the Mediterranean Basin.

the crocidurines, and more specifically that of Suncus, have been made through genetic approaches (Jenkins et al., 1998; Motokawa rouil et al., 2001; Dubey et al., 2007, 2008; Omar et al., 2000; Que et al., 2011; Meegaskumbura et al., 2012a, 2012b). According to some of these studies, S. etruscus diverged from other Eurasian Suncus and the remaining white-toothed shrews during the Late Miocene (7.2 Ma) (Dubey et al., 2008). Besides, several studies suggest that S. etruscus is genetically closer to Asian Suncus, whereas Afrotropical Suncus are closer to Sylvisorex (Dubey et al., 2007, 2008; Meegaskumbura et al., 2012a; Omar et al., 2011).

Indeed, Dubey et al. (2008) provide evidence that supports the idea that the genus Suncus is paraphyletic, with the Afrotropical Suncus species (S. aequatorius, S. lixa, S. hututsi, S. infinitesimus, S. remyi, S. varilla and S. megalurus) falling within Sylvisorex whereas S. etruscus belongs to a separate clade basal to the Asian Suncus and all crocidurine shrews. According to these results, afrotropical species of Suncus should be transferred to Sylvisorex, whereas the genus Suncus should be restricted to current Eurasian taxa.

Nevertheless, although in the updated Handbook of the Mammals of the World (Burgin and He, 2018) the paraphyly of Suncus is noted, the endemic African Suncus are still retained within the genus awaiting further research supporting this hypothesis. Therefore, in the light of these results an Asian origin of S. etruscus seems most probable.

The Etruscan shrew is the only species of the genus inhabiting both Europe and North Africa. Given the scarcity of its fossil record older than the Holocene, its provenance and palaeobiogeographical history is poorly known and, for this reason it has been subject of much debate (Heim de Balsac and Lamotte, 1957; Spitzenberger, 1990, Butler, 1998, Rzebik-Kowalska, 1998; Jenkins et al., 1998; rouil et al., 2001; Dubey et al., 2007, Motokawa et al., 2000; Que 2008; Omar et al., 2011; Meegaskumbura et al., 2012a, 2012b).

Spitzenberger (1990) suggested that this species might be an ancient European Tertiary relict component of the fauna considering its current broad geographic distribution and the existence of some European Pliocene records referred to the genus Suncus.

However, according to Rzebik-Kowalska (1998) all these records are taxonomically erroneous or at least, questionable.

Concerning the dispersal process into the Mediterranean Basin,

2. Small mammal assemblage of the Estrecho Cave The Estrecho Cave (Villares del Saz, Cuenca, Spain) is located at the central-eastern area of the Iberian Peninsula, on the western edge of Iberian System, more specifically at the unit of Sierra de Altomira (Fig. 1). The cave was formed in the Cretaceous limestones lez, 1998) and is part of what is (Diaz Molina and Lendínez Gonza known as the karst system of Villares del Saz. This system consists of a least four large cavities (the Monedas Cave, the Palomas Cave, the Camino Cave and the Estrecho Cave), some of them connected by long and narrow galleries. The Estrecho Cave entrance is found at an artificial near-vertical slope cut on a hill due to the construction of the A3 highway (Madrid-Valencia), close its 128 km.

These civil works left exposed a gallery of the karst system partially filled by sediments, leading thus to the discovery of the Estrecho

Fig. 1. Geographic location of the Estrecho Cave. Digital terrain model taken from Iberpix.

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Fig. 2. A. Plan of the Estrecho Cave (red area ¼ sampling area) modified from Ortega Martínez and Martín Merino (1992); B. Profile of the Estrecho Cave (red area ¼ sampling area) modified from Ortega Martinez and Martin Merino (1992); C. Stratigraphy of the sedimentary package of the entrance to the cave, with indication of the provenance of the small mammal assemblage level (CE-SE). (For interpretation of the references to colour in this figure legend, the reader is referred to the Web version of this article.)

Abundant microvertebrate remains have been recovered from a sedimentary package found at the entrance to the Estrecho Cave (Fig. 2). Preliminary data about the small mammals identified in this material was presented in Domínguez García et al. (2015).

Further information about the taphonomy and chronological features of this assemblage were described in Domínguez García et al.

(2019b), according to which the small mammal assemblage was originated by a mammalian predator of small to medium size.

A preliminary faunal list of the small mammal assemblage - the complete taxonomic study is currently in process - was included in Domínguez García et al. (2019b) in which 14 taxa belonging to four different mammal orders have been identified (Table 1). Concerning the chronology, a14C dating obtained from on a humerus of Arvicola sapidus gave a date of 2210 ± 15 BP, which has been recalibrated using OxCal v4.4.2 (Bronk Ramsey, 2009) and the new recent published calibration curve IntCal 20 (Reimer et al., 2020).

Thus, two intervals of calibrated dates were obtained: 2310e2290 (10.70%) and 2272e2149 (82,80%) years cal BP (Fig. 2c).

Cave (Ortega Martínez and Martín Merino, 1992).

In 1992 a prospecting campaign of the Villares del Saz karst system carried out by the Cultural and Sport Association “STD” and the Speleology Group “Edelweiss” resulted in an inventory of the cavities and of the Bronze Age archaeological remains found in the Estrecho Cave. Besides this, the finding of “a great accumulation of microvertebrate remains” in the uppermost part of the primitive entrance of the cave was reported (Ortega Martínez and Martín Merino, 1992).

In the year 2004 a new exploration was made by the Research Team “Lapis Specularis”, which confirmed the archaeological in mez terest of the cave and started research work in it (Bern ardez Go et al., 2005). In 2014 works to prepare the cave for touristic purpose rdez Go mez, 2016), triggering a started (Guisado di Monti and Berna multidisciplinary research project focused on the study of its geological features as well as its archaeological and palaeontological record.

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Table 1 Small mammals identified in the assemblage of the Estrecho Cave, taken from Domínguez García et al. (2019b) and unpublished data.

TAXON Rodentia Eliomys quercinus Arvicola sapidus Microtus cabrerae Microtus arvalis-agrestis Microtus duodecimcostatus-lusitanicus Apodemus sylvaticus-flavicollis Mus spretus-musculus Lagomorpha Oryctolagus cuniculus Eulipotyphla Crocidura russula Suncus etruscus Chiroptera Rhinolophus sp.

Myotis myotis-blythii Myotis sp.

Chiroptera indet.

2.

3.

3. Material and methods The specimens of the Etruscan shrew were found in samples collected in the Estrecho Cave in 2016. The material was taken from the sedimentary package located at the eastern edge of the entrance to the cave (Fig. 2a and b). In the uppermost part of this sedimentary package, a 50 cm thick detritic level with abundant microvertebrate remains was located (CE-SE). This level consists of a brownish-grey clay and limestone matrix with fragmented speleothems and centimetric clasts (Fig. 2c). In order to retrieve the small-vertebrate remains a water-screening system with two superimposed sieves of decreasing mesh size (2 and 0.5 mm) was used.

The taxonomic identification of the Suncus material was based on the morphometric criteria described in specialized literature (Chaline et al., 1974; Gos albez, 1987; Blanco, 1998; Sanz Navarro n Artigas, 2017). Additionally, the fossil remains were and Turo compared with recent Etruscan shrew skeletons from C. Laplana’s personal collection. Nomenclature and measurements follow SansComa et al. (1981). Measurements employed in the biometric analysis are available as supplementary material. Taxonomy has been standardized according to Burgin and He (2018), in which Crocidura suaveolens group systematics has changed. Therefore, populations from Western and Central Europe, Mediterranean islands and southwestern Asia are considered as Crocidura gueldenstaedtii, while Crocidura suaveolens is retained for East European and Asian populations.

In order to reconstruct the biogeographic history of the Etruscan shrew in the circum-Mediterranean region, a detailed critical review of published records of this species was conducted. To analyse the reliability of these records, the taxonomic precision and dating methods associated to these data were considered critically.

4.

5.

as we move from Neolithic to Protohistoric and Historic times (Papayiannis, 2012; Laplana and Sevilla, 2013). This is primarily due to the lower value assigned to small mammals as tools for chronological and environmental interpretations of Late Holocene archaeological sites compared to that in Pleistocene and Early Holocene sites. For this reason, there is an important drop in available data in comparison to those from the Late Pleistocene and Early Holocene, having consequences on other limitations commented further on.

Collecting methods are commonly inadequate to obtain a good representation of the small mammals present in the assemblages. It is a common procedure in Holocene sites to dry-sieve the sediments and only in few sites water-screening systems with a mesh below 2 mm are used; consequently, retrieval of small vertebrates is severely biased (Cucchi et al., 2005; Domínguez García et al., 2019a). This is particularly relevant for small vertebrates, and considering the tiny skeletal elements of S. etruscus, it can be taken for certain that its known fossil record underestimates its representation in the past (Arribas, 2004).

Etruscan shrew records are often inaccurate from a taxonomic point of view. Despite its distinctly smaller size compared to Crocidura spp., or the presence of four instead of three antemolar unicuspid teeth on the maxilla, verifiable records are scarce, as they are commonly lacking descriptions, morphometric data and figures of the fossils.

The imprecise age of the small vertebrate assemblages. In many Holocene sites, the age of the associated small mammal assemblages is rather imprecise, since direct absolute dating is rarely performed. The small mammal assemblages are assigned an age according to the archaeological context, under the assumption they are contemporaneous, which is not always so.

The stratigraphy in these sites is frequently altered by recent bioturbation, implying faunal mixing with the intrusion of modern remains into older materials.

S. etruscus skeletal remains are neither abundant in recent pellets or scats, nor in fossil small mammal assemblages. The Etruscan shrew is a common prey for many predators of small mammal, such as barn owls, short-eared owls, long-eared owls, little owls, genets or beech martens. However, it is known to represent a low percentage in their diet compared to other prey pez-Fuster et al., 1979; Lo pez-Fuster, 2007; Guillem (Lo Calatayud, 1996; Torre et al., 2013; Domínguez García and Gamboa, 2017). Having in mind that the main origin of fossil microvertebrate assemblages is predation (Andrews, 1990; ndez-Jalvo et al., 2016), a low representation of the Ferna Etruscan shrew in fossil sites is not surprising.

5. Results After processing 150 Kg of sediment from the Estrecho Cave, three remains belonging to S. etruscus - two mandibles and an upper second molar (M2) e were recovered. They were assigned to the pigmy white-toothed shrew on the basis of morphology and size (Figs. 3e6). Three white-toothed shrews (subfamily Crocidurinae) may potentially be present in this assemblage (Crocidura russula, Crocidura gueldenstaedtii and Suncus etruscus). The Etruscan shrew is distinctly smaller than Crocidura spp., besides having four upper antemolar unicuspid teeth, whereas Crocidura spp. have only lbez, 1987; Blanco, 1998). Unfortuthree (Chaline et al., 1974; Gosa nately, since no maxillae were found, this character could not be used here. However, the difference in size between S. etruscus and Crocidura was decisive, the remains from Estrecho Cave falling well within S. etruscus range and outside the range displayed in

4. Data limitations Five main issues that limited our palaeobiogeographical analysis were derived from the characteristics of the archaeopalaeontological record of S. etruscus: 1. Amount of interest focused on the small mammals in Holocene archaeo-palaeontological sites. The number of specialized studies focused on small mammal fossil assemblages decreases 4

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Aven Marzal II in Mediterranean France (Debard et al., 1999), the Late Pleistocene levels from the Noisetier Cave in France (Jeannet, 2001), the Ibex Cave in Gibraltar (Denys, 2000) and the Cueva Nueva in Spain (Barea et al., 2002). However, the Pleistocene records from Western Europe are less reliable than those coming from the Eastern Mediterranean. In particular, the Qesem Cave records of S. etruscus are documented with morphometric data and figures (Maul et al., 2011), and the age is well established by TL/and ESR/U-series dating methods (Mercier et al., 2013). On the other hand, the record from Pretalona must be considered with caution as it consists of a single lower incisor (Rzebik-Kowalska, 1998).

As mentioned, Pleistocene records from Western Mediterranean Europe are highly doubtful. The earliest record would be that from Aven Flahaut or Aven Marzal II (France), where the Etruscan shrew is mentioned in the assemblage 1 (Debard et al., 1999). Regarding the chronology of this record, the upper part of this section was dated by 14C as Late Pleistocene, whereas the intermediate levels (assemblages 2 and 3) were dated by an analysis of the composition of tephra found in the filling sediments around 450 ka BP (Debard and Pastre, 2008). According to the small mammals, lower levels (1 and 2) were dated as Middle Pleistocene based on the presence of Pliomys lenki (Debard et al., 1999). However, this species is known to have a longer record since it is known from several sites of latest pez-García, 2011) as Pleistocene age from the North of Spain (Lo well as in Early Holocene sites from France (Marquet, 2001).

Therefore, this assemblage could be in fact younger than reported.

Furthermore, this mention is taxonomically unreliable, no description of the material nor figures are included in the paper to justify the identification. Accordingly, it is necessary to carry out a review of these small mammal assemblages along this sequence to confirm o dismiss both the S. etruscus identification and their chronology. The second oldest record in Western Europe would be that coming from Unit 3 from Ibex Cave dated as Late Pleistocene (49e37 ka BP) according to Barton et al. (1999) and Denys (2000).

Unfortunately, these remains were published as “Crocidurinae indet. (cf. Suncus)”, and therefore taxonomically imprecise. Another Late Pleistocene record of S. etruscus comes from Level 1 of Noisetier Cave dated from 42e33 ka BP (Jeannet, 2001; Mourre et al., 2008). However, in this site the Etruscan shrew was only found in the uppermost sublevel of Level 1, which is immediately below the top level and could be younger or undergone recent contamination.

Besides, this record is again taxonomically imprecise, since measurements, descriptions or figures of the material are not provided.

The latest Pleistocene record found is mentioned by Barea et al.

(2002) in a preliminary study that reports the finding of small vertebrate remains at the Cueva Nueva in Pedraza, Spain. This occurrence is once again taxonomically imprecise since it is mentioned as “Soricidae gen. indet. cf. Suncus sp.” and no descriptions or figures are provided. Concerning its age, it is undetermined since no dating has been performed and there are no archaeological remains associated to this material. The authors proposed a maximum age of Late Pleistocene on the basis of the presence of Rattus cf. rattus in the assemblage. However, the presence of this species indicate a considerably younger age for the assemblage since this taxon it does not appear in the Iberian Peninsula until the second half of the third millennium BP (AudoinRouzeau and Vigne, 1994; Domínguez García et al., 2019a). In more recent chronologies, specifically in the Holocene, the number of Etruscan shrew occurrences increases substantially. Thus, it has been mentioned in Turkey (Jenkins, 2012), Azerbaijan (Berthon et al., 2013), Jordan (Pokines and Ames, 2015), Iberoccitanian Region (Asquerino and Woloszyn, 1991; Jeannet and Vital, 2009; Poitevin et al., 2005, this work) and, with the exception of the Balearics, it has been found in several Mediterranean islands such

Fig. 3. Suncus etruscus remains from the Estrecho Cave. A) Right mandible, labial view (CE-SE-41); B) Left mandible, labial view (CE-SE-61); C) Right M2, occlusal view (CESE-42). Scale bars: 1 mm.

Crocidura spp. The height of coronoid process (HC) is particularly relevant to differentiate between both possibilities, which gives a value for this parameter going from 2.5 to 3 mm in S. etruscus from the Iberoccitanian Region, while in Crocidura spp. attain values over n 3.5 mm (Chaline et al., 1974; Blanco, 1998; Sanz Navarro and Turo Artigas, 2017). This parameter mesaured on the single mandible retaining the coronoid process in the Estrecho Cave (Fig. 3a; Fig. 4) gave a value for HC of 2.805 mm. Additional biometric analyses performed on the three remains showed that this material from the Estrecho Cave is distinctly smaller than Crocidura spp. and their values fall within the range of variation of S. etruscus from both modern and fossil localities for all the variables (Figs. 4e6).

5.1. Palaeobiogeographical review The fossil record of the Etruscan shrew is rather scarce, probably enhanced by the limitations mentioned above. Nevertheless, a total number of 28 records of S. etruscus were found in the Mediterranean Basin (Table 2, Fig. 7). The oldest records come mainly from Middle and Late Pleistocene sites of Israel (Tchernov, 1996, 1998; Maul et al., 2011; Comay et al., 2019). Besides, there are another five Pleistocene records which come from European sites: the Greek Middle Pleistocene cave of Petralona (Pachyura cf. etruscus) (Sickenberg, 1971), the Middle Pleistocene site of Aven Flahaut or 5

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Fig. 4. Height of coronoid process (HC) from the Estrecho Cave in comparison with recent localities and Holocene sites with S. etruscus, C. gueldenstaedtii and C. russula from n (SP); 3: Catalonia (SP); 4: S. France; 5: S. France Mediterranean region; bars display mean, minimum and maximum values. 1: Estrecho Cave (CE-SE-41); 2: Fuentes de Leo (pellets); 6: Huesca-Pyrenees (SP); 7: SE. Bulgaria; 8: SE. Anatolia (TU); 9: Italy; 10: Iran; 11: Portugal; 12: Korfu (GR); 13: Kos (GR); 14: Turkey; 15: Chios (GR); 16: Kouklia; 17: La res-Sonneville (FR); 19: Basque Country (SP); 20: Linares de Riofrío (SP); 21: Tharandt (GER); 22: Fülo €ph aza (HUN); 23: Lucca (IT); 24: Monte Gargano (IT); Algaida (SP); 18: Lingnie n (SP); 31: Burgos (SP). Sources of data: 1: This work; 2: Domínguez 25: La Algaida (SP); 26: Bonn (GER); 27: Ramales de la Victoria (SP); 28: Portugal; 29: Asturias (SP); 30: Leo pez-Fuster et al. (1979); 4, 5: Sans-Coma et al. (1981); 6, Vericad (1971); 7: Popov et al. (2004); 8: Coşkun and Kaya (2013); 9: Contoli et al. (2000); García and Gamboa (2017); 3: Lo 10: Esmaeili et al. (2008); 11, 12, 28: Niethammer (1970); 13, 14, 21e24: Niethammer and Krapp (1990); 15: Besenecker et al. (1972); 16: Reumer and Oberli (1988); 17, 25: Rey and Landin (1973); 18: Mottaz (1908); 29, 30, 31: Zabala (1985); 20: Vesmanis and Kahman (1976); 26: Sans-Coma and Margalef (1981); 27: Niethammer (1964); 29: García-Dory (1977).

Abbreviations: SP: Spain; TU: Turkey; GR: Greece; FR: France; GER: Germany; HUN: Hungary; IT: Italy; SE: Southeast.

of S. etruscus from the Early Holocene are known from the “Rattus rattus layer” of Ghar Dalam cave in Malta (Storch, 1974).

Regarding the Western Mediterranean Islands, two sites are known to have yielded remains of S. etruscus. First, the Middle Holocene record found at Su Guanu Cave in Sardinia (Sanges and Alcover, 1980; Sans-Coma et al., 1985), with both imprecise dating and unreliable stratigraphy. The small mammal assemblage containing the Suncus remains is supposed to be older than human occupations of the cave (radiocarbon dated in 5742e5489 cal BP).

However, this assemblage also yielded Rattus rattus and Mus musculus remains, which are known to be absent from the western Mediterranean area prior to 3000 years BP (Audoin-Rouzeau and Vigne, 1994; Vigne and Valladas, 1996; Cucchi et al., 2005). Thus, the presence of these two latter species may indicate a younger age than previously supposed for this record. The second record comes from the Late Holocene site of Monte di Tuda in Corsica, where a rich small mammal assemblage has been studied (Vigne and Valladas, 1996). As a result, three well dated biozones were distinguished in its stratigraphic sequence, in which S. etruscus is well documented in all three, from the oldest (Biozone 3; 2814e2101 cal BP) to the youngest (Biozone 1; after 726-491 cal BP).

In Southwestern Europe mainland (Iberian Peninsula and Mediterranean France) four sites of Holocene age have yielded

as Chios (Besenecker et al., 1972), Crete (Reumer and Payne, 1986; Papayiannis, 2012); Cyprus (Reumer and Oberli, 1988); Malta pez-García et al., (Storch, 1974; Hunt and Schembri, 1999); Sicily (Lo 2013); Sardinia (Sanges and Alcover, 1980; Sans-Coma et al., 1985) and Corsica (Vigne and Valladas, 1996) (Table 2, Fig. 7).

In the Eastern Mediterranean, most of the records can be considered taxonomically reliable since they are accompanied by taxonomic descriptions and measurements of the material, and were collected in well documented archaeological contexts (Besenecker et al., 1972; Papayiannis, 2012; Reumer and Payne, 1986; Reumer and Oberli, 1988). Most of them belong to Late Holocene sites (Minoan Crete and Late Bronze Age of Cyprus), with the exception of the record from the Aegean island of Chios (Greece), which unfortunately has not been dated, although it is most probably of Early Holocene age according to Besenecker et al.

(1972).

In the central Mediterranean, at Vallone Inferno rock-shelter (Sicily) a fragmented mandible of Etruscan shrew was found in level 3.4, which unfortunately has been assigned a relatively wide chronological interval, due to radiocarbon age results of 4500 to pez García et al., 2013). Again, although the authors 3360 cal BP (Lo mention “the tiny length dimensions of the molar series make this record unambiguous”, no measurements nor figures are given in the paper, so this record cannot be confirmed. More recent records 6

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sites. One of them is the Cueva de los M armoles in Spain (Asquerino and Woloszyn, 1991); the other is the Chauve-Souris Cave in France (Jeannet and Vital, 2009). In both cases these records must be considered uncertain due to the imprecise taxonomic identifications and to the inaccurate dating of the assemblages (Domínguez García et al., 2019a). For the Late Holocene, besides the record from the Estrecho Cave described in this work, S. etruscus has also been described at Puits de Lattara in France, where the species is well represented in a Roman context (Poitevin et al., 2005).

6. Discussion Given the available data, the accuracy of S. etruscus fossil record should be treated with caution, since the taxonomy and chronology of only 9 of the 28 occurrences are fully documented. Nevertheless, these data provide a relatively clear picture of the route, timing and mechanisms involved in the dispersal process of the Etruscan shrew in the Mediterranean Basin.

1. S. etruscus was present at least in the Middle Pleistocene of Israel, and it appears restricted to the Levant in Eastern Mediterranean during the whole Pleistocene. There are few European Pleistocene records, but of low reliability. Even though these records might indicate an introduction into Europe earlier than the Holocene, they would not represent a successful colonisation persisting until the present, since it is highly unlikely that the Etruscan shrew could have survived in the harsh climatic conditions of glacial periods in Europe, particularly in the cases of Pleistocene records from France, given that this species shows pez-Fuster, 2007; Burgin Mediterranean climatic affinities (Lo and He, 2018). Indeed, repeated changes of mammal geographic ranges during the Pleistocene have been recognised (Schreve, 2019). Nevertheless, leaving aside these doubtful occurrences, the numerous updated and well documented works of synthesis of the rich and well-known Pleistocene small mammal record of the Iberian Peninsula, France, Italy, Croatia and North Africa, in which absolutely no S. etruscus remains have been found (Poitevin et al., 1990; Vigne and Pascal, 2003; pez-García, 2011; Lenardi c et al., 2018; Stoetzel, 2013; Stoetzel Lo et al., 2019; Berto et al., 2019), support the idea that the Etruscan shrew was absent from these areas during the whole Pleistocene.

All this data therefore, support the hypothesis of an Asian origin of the Etruscan shrew instead of an African origin pro rouil posed by several authors (Rzebik-Kowalska, 1998; Que et al., 2018). They also et al., 2001; Garrido-García, 2008; Furio indicate that successful colonisation of Europe and North Africa did not take place until the Holocene. Thus, we can state that the dispersal process of the Etruscan shrew through the Mediterranean Basin took place westward reaching the Iberian Peninsula at the latest in the Late Holocene.

2. Another point of interest is the process by which this species was introduced into some Mediterranean islands. Given that during the Holocene most large Mediterranean islands were completely separated from the mainland, a natural introduction of S. etruscus into these islands through natural dispersal processes can be discarded. Its anthropophilous nature (Bergier, 1988; Vigne, 1999; Aulagnier et al., 2017) may have favoured the accidental introduction of the species into the Mediterranean islands. Therefore, it seems most likely that accidental anthropic translocations of the species took place during the Holocene along navigation routes, probably transported as stowaways within infested grain and foodstuffs cargo (Dobson, 1998; Vigne, 1999, 2014; Garrido-García, 2008; Omar et al.,

Fig. 5. Length from the first to the third lower molar (Lm1-m3) from the Estrecho Cave in comparison with recent localities, Pleistocene and Holocene sites of S. etruscus, C. gueldenstaedtii and C. russula from Mediterranean region; bars display mean, minimum and maximum values. 1: Estrecho Cave (CE-SE-41, CE-SE-61); 2: Catalonia (SP); 3: S. France; 4: S. France (pellets); 5: SE. Bulgaria; 6: Crete (GR); 7: Kouklia; 8: Kom€ ph aza (HUN); 12: Lucca (IT); 13: mos; 9: Quesem Cave; 10: Tharandt (GER); 11: Fülo pez-Fuster Monte Gargano (IT); 14: Morges (SWI). Sources of data: 1: This work; 2: Lo et al. (1979); 3, 4: Sans-Coma et al. (1981); 5: Popov et al. (2004); 6, 8: Reumer and Payne (1986); 7: Reumer and Oberli (1988); 9: Maul et al. (2011); 10e14: Niethammer and Krapp (1990). Abbreviations: SP: Spain; GR: Greece; GER: Germany; HUN: Hungary; IT: Italy; SWI: Switzerland; SE: Southeast.

Fig. 6. Bivariate plot of the length of the second upper molar (LM2) vs. width of the second upper molar (WM2) from the Estrecho Cave in comparison with recent localities, Pleistocene and Holocene sites with S. etruscus, C. gueldenstaedtii and C. russula n (SP); 3: from Mediterranean region. 1: Estrecho Cave (CE-SE-42); 2: Fuentes de Leo Catalonia (SP); 4: S. France; 5: S. France (pellets); 6: SE. Bulgaria; 7: Cova de les Cendres (SP); 10: PRERESA (SP); 11: (SP); 8: Cova Colomera (SP); 9: Castellet del Bernabe Albacete (SP); 12: Kouklia; 13: Cyprus; 14: Kommos; 15: Crete. Sources of data: 1: This pez-Fuster et al. (1979); 4, 5: Sans-Coma et al. (1981); 6: work; 2: This work; 3: Lo pez-García (2011); 9: Gue rin Popov et al. (2004); 7: Guillem Calatayud (2009); 8: Lo et al. (2011); 12, 13: Reumer and Oberli (1988); 14, 15: et al. (1989); 10, 11: Sese Reumer and Payne (1986). Abbreviations: SP: Spain; SE: Southeast.

remains of S. etruscus. Starting with the Middle Holocene related to Neolithic cultural contexts, S. etruscus has been mentioned in two 7

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Table 2 Suncus etruscus records in the Mediterranean Basin. References: 1: Ferrandi et al. (1994); 2: Pokines and Ames (2015); 3: Poitevin et al., (2005); 4: This work; 5: Storch (1974), pez-García et al. (2013); 12. Jeannet and Vital (2009); 6: Vigne and Valladas (1996); 7: Reumer and Oberli (1988); 8: Reumer and Payne (1986); 9, 10: Papayiannis (2012); 11: Lo 13: Sanges and Alcover (1980), Sans-Coma et al. (1985); 14: Berthon et al. (2013); 15: Asquerino and Woloszyn (1991); 16: Besenecker et al. (1972); 17: Jenkins (2012), Cessford (2001, 2005); 18: Barea et al. (2002); 19: Comay et al. (2019); 20: Jeannet (2001); 21: Denys (2000); 21, 22, 23, 25: Tchernov (1998); 25: Mercier et al., 2007; 26: Maul et al.

(2011); 27: Sickenberg (1971); 28: Debard et al. (1999). Abbreviations: ka: thousands of years, FR: France, SP: Spain, GR: Greece, IT: Italy, UK: United Kingdom, IS: Israel.

Time interval

Nº

Chronology

Site/Level

Locality/Country

Late Holocene

1 2 3 4 5 6 7 8 9 10 11 12 13

Oletta Cave (Level D) Wadi Zarqu Ma’in 1 Puits de Lattara Estrecho Cave Ghar Dalam (Levels I-III) Monte di Tuda Kouklia Kommos Khania Mochlos Vallone Inferno (Level 3.4) Chauve-Souris Cave (Level 15) Su Guanu Cave

Corsica/FR Jordan Lattes/FR Villares del Saz/SP Malta Corsica/FR Cyprus Crete/GR Crete/GR Crete/GR Sicily/IT re/FR Donze Sardinia/FR

14 15

1052e548 cal BP 1290e1170 cal BP 2050 BP- 1800 BP 2311e2154 cal BP 2700 BP-present 2814-491 cal BP 3300e3200 BP 3320e3150 BP 3325e3150 BP 3470 BP 4500e3360 cal BP 4814e4086 cal BP >5742e5489 cal BP 5660e5535 cal BP 6300e6200 cal BP ?

Azerbaijan rdoba/SP Priego de Co

16 17 18 19 20 21 22 23 24 25 26 27 28

?

9350e7950 cal BP ?

34-33 ka BP 42-33 ka BP 49-37 ka BP 61-45 ka BP 105-85 ka BP 115- 92 ka BP 225-115 ka BP 280-250 ka BP ? >450 ka BP

Ovçular Tepesi (Pit 01.171) Cueva de los rmoles (Area F) Ma Chios €yük Catalho Cueva Nueva Manot Cave Noisetier Cave (Level 1) Ibex Cave (Unit 3) Kebara (VI-XII) Tabun B Qafzeh (XV-XXV) Hayonim (Lower E) Qesem cave (Concentration 1) Petralona Cave Aven Flahaut/Marzal II (Assemblage 1)

Middle Holocene

Early Holocene Late Pleistocene

Middle Pleistocene

Greece Turkey Spain Israel chet/FR Fre Gibraltar/UK Mount Carmel/IS Mount Carmel/IS Nazaret/IS Galilee/IS Israel Greece che/FR Arde

Fig. 7. Map of Suncus etruscus occurrences in the Mediterranean Basin. Numbers used for each occurrence refer to the sites listed in Table 2. Modified from CC-BY 4.0 ign.es 2015.

into Anatolia and into Central Asia. However, more reliable records are necessary to confirm this interpretation.

4. Concerning the timing of the Holocene dispersal process of S. etruscus across Mediterranean Sea, the occurrence from Chios (Besenecker et al., 1972) is so far the oldest known record. Unfortunately, it cannot be taken for certain due to the imprecise chronology assigned to this assemblage. Some Middle Holocene pez García et al., 2013) and records found in Central (Sicily, Lo Western Mediterranean Islands (Sardinia, Sanges and Alcover, 1980; Sans-Coma et al., 1985) seem to support an earlier introduction of S. etruscus in this area than in the Eastern Mediterranean islands, since the arrival of the Etruscan shrew to Crete

2011). Indeed, the S. etruscus remains studied by Reumer and Oberli (1988) from Kouklia (Cyprus) were found in the contents of a storage vessel. Direct evidence of stowaway transport of small mammals in the Mediterranean Basin has been described in the Late Bronze Age Uluburun shipwreck, where Mus musculus domesticus was identified (Cucchi, 2008). Therefore, our results are in accordance with previous hypothesis for a human-facilitated dispersal process of small mammal species in the Mediterranean Basin during the Holocene.

3. The Early Holocene records found in Turkey (Jenkins, 2012) and Middle Holocene from Azerbaijan (Berthon et al., 2013) might evidence the first steps of terrestrial spreading of this species 8

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place after several arrival events that enable stability and growth of pioneering populations (Blondel, 1986; Blondel and Vigne, 1993), the intensification of seafaring might have also increased the numbers of migrant S. etruscus in westward destinations, favouring its successful introduction in larger Mediterranean islands and in southwestern European mainland.

Furthermore, according to Vigne (1999, 2014) and Vigne et al.

(2014) the appearance of new types of boats with decks could also have facilitated the transport of small mammals hidden within them. Seemingly, the dispersal process of S. etruscus into the Mediterranean followed the same route and took place at about the same time as that of other small mammal species such as Mus musculus domesticus and Rattus rattus (Audoin-Rouzeau and Vigne, 1994; Cucchi et al., 2005).

However, it must be borne in mind that the age inferred for the spreading of S. etruscus along the Mediterranean Basin through this analysis is only tentative due to the scarcity of reliable records, some of which are of doubtful age. For instance, the time gap between the first occurrence of Mus musculus domesticus in Cyprus (11,000e10,000 cal. BP) and that of S. etruscus (3300e3200 BP) is striking since seafaring from Cyrpus to its near mainland was intensive since the beginning of the Holocene (Cucchi et al., 2002, 2005; Cucchi, 2005; Vigne et al., 2014). An earlier translocation cannot be ruled out, taphonomic or methodological reasons being behind the absence of S. etruscus until a later date. Alternatively, a later colonisation of S. etruscus is also reasonable if we consider that the introduction of M. musculus domesticus was probably favoured by its commensal nature.

and Cyprus is well documented and dated as having occurred not earlier than the Late Holocene, around the mid-fourth millennium BP (Reumer and Payne, 1986; Reumer and Oberli, 1988; Reumer, 1996; Papayiannis, 2012). On the other hand, its introduction in Malta (Storch, 1974), and Corsica (Vigne and Valladas, 1996) apparently took place later, as the earliest records of the species in these islands is dated as the beginning of third millennium BP. However, this apparent anomaly in the westward spreading of the species may simply be a consequence of the age assigned to Sardinian and Sicilian records, which are not fully precise. Thus, in overall the available reliable records seem to support a scenario in which the Etruscan shrew gradually spread through the Mediterranean Basin from the East to the West, starting approximately 3500 years ago.

It is unfortunate that the known palaeontological record of the Etruscan shrew in European mainland has very few reliable records. The available data show that the arrival and colonisation of Mediterranean France and Iberian Peninsula probably had already taken place by the end of the third millennium BP. Leaving out some doubtful records of earlier age, the occurrence from the Estrecho Cave is so far the most reliable record of S. etruscus in European mainland. The lack of records of this species among the small mammal remains retrieved in Iberian older sites where meticulous collecting methods were used, obtaining rich, diverse and well dated assemblages such as in El ~ uls Cardona et al., 2017), Castillejo del Bonete Mirador (Ban (Domínguez García et al., 2019a) or several sites from the Spanish Levant (Guillem Calatayud, 1999), point to the absence of S. etruscus in the Iberian fauna until well into the Late Holocene.

5. Regarding the highly likely relationship between the development of navigation techniques and the dispersal process of S. etruscus along the Mediterranean, although there are indirect evidences of seafaring even from the Middle Pleistocene and the Late Pleistocene, these are not frequent and mainly related to small-scale and cabotage trips (Bednarik, 2003; Broodbank, 2006; Abulafaia 2011; Zilh~ ao, 2014; Howitt-Marshall and Runnels, 2016; Papoulia, 2016, 2017; Moncel et al., 2020), and therefore it is unlikely that translocations took place by this means at that time. However, the exchanges and sea traffic increased during the Holocene from around 10,000 to 7300 BP, related to the East to West Neolithic expansion (Cucchi et al., ~o, 2014; de Vareilles 2005; Zeder, 2008; Abulafaia 2011; Zilha et al., 2020). This process involved the establishment of farming communities around the Mediterranean Basin, and involved translocations of both domesticated and wild animals, as well as plants (Cherry, 1990; Audoin-Rozeau and Vigne, 1994; Vigne, 1999; 2014; Cucchi et al., 2005; Zeder, 2008; Domínguez García et al., 2019a; de Vareilles et al., 2020). However, these contacts between eastern and western Mediterranean shores were still geographically limited and it is not until the beginning of the third millennium BP that the Mediterranean Sea became widely crossed by frequent East-West seafaring, mainly related to the commercial activities of Phoenicians, Greeks and later on, the Roman Empire (Cucchi et al., 2005; Cucchi and Vigne, 2006; ~o, 2014; Domínguez García et al., 2019a).

Abulafia, 2011; Zilha This notable growth in displacements can also be detected from genetic studies of Iberian human populations. These indicate an initial migration founder bottleneck in early farmer Neolithic populations arriving from the Eastern Mediterranean, followed by admixture with local previous populations (Valdiosera et al., 2018), but with the main genetic transformation occurring by gene flow from the Eastern Mediterranean and North Africa as far back as the Roman period (Olalde et al., 2019).

Given that successful colonisation by a species only takes

7. Conclusions As concluding remarks, in agreement with what several authors have also pointed out (Cucchi et al., 2005; Stewart, 2005; Bell et al., 2010), we would like to draw attention on the importance of having a precise and well dated fossil record to reconstruct small mammal dispersal events during the Pleistocene and Holocene. Thus, as exposed here, scanty material and doubtful chronology limits at present the precise reconstruction of how and when the Etruscan shrew was introduced in Europe. The few well dated and well identified fossil material of this species, such as the record from the Estrecho Cave acquires for this reason special relevance. Although some aspects must still be addressed, such as the time in which S. etruscus arrived to the Balkans, Italian Peninsula and to the Maghreb, all reliable data seem to point towards a similar pattern of colonisation as that described for M. musculus domesticus and Rattus rattus, which could be a human-facilitated dispersal process from eastern to western Mediterranean, beginning around the middle of the 4th millennium BP. However, further research is needed to confirm and reinforce this hypothesis. Therefore, new material and the review of material coming from Holocene sites with small mammal assemblages could possibly increase its records and help answer this question.

Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Acknowledgments We would like to thank the team Lapis Specularis, specially to 9

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Bern mez, for Juan Carlos Guisado di Monti and María Jose ardez Go allowing us to carry out the sampling of small mammal deposits within their Archaeological Intervention Project for the adaptation of the Estrecho Cave for tourist use (Expte:15.1441 of the Government of Castilla la Mancha). This work is a contribution to the projects PGC2018-094125-B-100 (MCIU/AEI/FEDER, UE) and CGL2016-79334-P of the Spanish Ministry of Science, Innovation and Universities. This is a contribution by the Research Group UCM 970827 on Quaternary Ecosystems of Complutense University of Madrid. The first author, ACDG, has a predoctoral grant from the Complutense University of Madrid (CT17/17-CT18/17).

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Domínguez García, A.C., Laplana, C., Sevilla, P., Blain, H.A., Palomares Zumajo, N., Benítez de Lugo Enrich, L., 2019a. New data on the introduction and dispersal process of small mammals in Southwestern Europe during the Holocene: Castillejo del Bonete site (Southeastern Spain). Quat. Sci. Rev. 225, 106008. https://doi.org/10.1016/j.quascirev.2019.106008. rdez Domínguez García, A.C., Laplana, C., Sevilla, P., Guisado di Monti, J.C., Berna mez, M.J., 2019b. Tafonomía y cronología de la asociacio n de micromamíferos Go de la Cueva del Estrecho (Villares del Saz, Cuenca). Rev. Esp. Paleontol. 34 (2), 241e256 https://doi.10.7203/sjp.34.2.16114. re, P., Vogel, P., 2007. Molecular phyloDubey, S., Salamin, N., Ohdachi, S.D., Barrie genetics of Soricidae (Mammalia, Eulipotyphla) reveals timing of transcontinental colonizations. Mol. Phylogenet. Evol. 44, 126e137. https://doi.org/ 10.1016/j.ympev.2006.12.002. re, P., Colyn, M., Vogel, P., 2008.

Dubey, S., Salamin, N., Ruedi, M., Barrie

Appendix A. Supplementary data Supplementary data to this article can be found online at https://doi.org/10.1016/j.quascirev.2020.106690.

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