Quaternary International 389 (2015) 119e130 Contents lists available at ScienceDirect Quaternary International journal homepage: www.elsevier.com/locate/quaint How rare was human presence in Europe during the Early Pleistocene? * Jesús Rodríguez a, , Ana Mateos a, Jesús Angel Martín-Gonzalez b, 1, Guillermo Rodríguez-Gomez a a CENIEH, Paseo Sierra de Atapuerca, 3, 09002 Burgos, Spain b Dpt. Matematicas y Computacion, Universidad de Burgos, Burgos, Spain article info abstract Article history: Beneath the hot debate about the tempo and mode of the first human colonization of Europe is the Available online 30 December 2014 perception that the record of human presence in the Early Pleistocene is sparse and fragmented. As a result, it is often implicitly assumed that hominins, if present, were scarce in the Early Pleistocene Eu- Keywords: ropean ecosystems. Here we present a quantitative assessment of the rarity and commonness of the Rarity European large mammal species during the 1.4e0.8 Ma period, including hominins. Considering the Hominin palaeontological record only, Homo was not one of the most common species in Europe, but it may not be Early settlement considered a rare species. In contrast, taking into consideration the archaeological record, hominins Geographical range Occupancy exhibit a wide geographical distribution and a high frequency of occurrence (occupancy) in comparison with other large mammals. It is speculated that hominins were frequent but not abundant in Europe during the late Early Pleistocene. © 2015 Elsevier Ltd and INQUA. All rights reserved. 1. Introduction Mosquera et al., 2013; Martínez et al., 2014). Some theoretical evolutionary scenarios have been proposed including argumen- Timing and nature of the early human colonisation of Europe tations in pro of a continuous settlement (Bermúdez de Castro and has been a hotly debated topic for a long time. Nowadays, there is Martinon-Torres, 2013; Garcia et al., 2013; Martínez et al., 2013) a general consensus in assuming that humans dispersed to and/or supporting a discontinuous and intermittent occupation by Western Europe during the late Early Pleistocene (Carbonell et al., populations constrained by climatic and ecological fluctuations 2010; Made and Mateos, 2010; Moncel, 2010; Palombo, 2010, (Dennell, 2003; Agustí et al., 2009; Dennell et al., 2011; Leroy et al., 2013; Allue et al., 2013). It is generally accepted that humans 2011; MacDonald et al., 2012). These debates reflected the usual occupied Europe as early as 1.4 Ma as evidenced by the fossils and problems derived from the low completeness of the hominin fossil lithic industry recovered at Barranco Leon D (Spain) and Pirro record, which represents a fragmented, poorly-dated set of evi- Nord (Italy) (Arzarello et al., 2007; but see Muttoni et al., 2013, dences. Several issues related to the ecological and environmental 2011; but see Rolland, 2013; Toro-Moyano et al., 2013). Beyond context of this dispersal event, or the technological skills of these the debate about the phylogenetic relationships between those populations are still under discussion (Turner, 1992; Arribas and hominin European populations (Stringer, 2012; Gomez-Robles, Palmqvist, 1999; Mosquera et al., 2013). Moreover, recent discov- 2013; Lordkipanidze et al., 2013; Arsuaga et al., 2014), a new eries and critical evaluations of the chronological framework and debate aroused in the last years concerning the continuity or dispersal events related to the first human colonization of Europe discontinuity of this initial settlement (Dennell, 2003; Leroy et al., (Muttoni et al., 2011, 2013; Pares et al., 2013; Rolland, 2013) show 2011; Muttoni et al., 2011; Bermúdez de Castro et al., 2013; that key questions about the tempo and mode of this colonization remain unsolved. Several models have been proposed supporting the idea that * Corresponding author. the first human colonisers were restricted to the Mediterranean E-mail addresses: [email protected] (J. Rodríguez), ana.mateos@cenieh. area because they were unable to survive in mid-latitude Europe es (A. Mateos), [email protected] (J.A. Martín-Gonzalez), guillermo. (Roebroeks and Kolfschoten, 1992; Dennell and Roebroeks, 1996; [email protected] (G. Rodríguez-Gomez). 1 Temporarily assigned to CENIEH. Roebroeks, 2001; Dennell et al., 2010). Although many of the http://dx.doi.org/10.1016/j.quaint.2014.12.016 1040-6182/© 2015 Elsevier Ltd and INQUA. All rights reserved. 120 J. Rodríguez et al. / Quaternary International 389 (2015) 119e130 evolutionary scenarios proposed take into account only the highly constrained opportunities of the settlement for early palaeoanthropological and/or archaeological evidence (Carbonell hominins. et al., 1999; Dennell et al., 2010, 2011; MacDonald et al., 2012; All things considered, human occupation during the Early Barsky et al., 2013; Bermúdez de Castro and Martinon-Torres, Pleistocene is currently represented in Europe by a short list of sites. 2013; Mosquera et al., 2013), it becomes increasingly evident Many authors assume that the fragmentary archaeological and that some complex relationships existed between palaeoclimatic fossil records indicate only sporadic hominin presence or marginal and palaeoecological changes and human dispersals and settling settlement (MacDonald et al., 2012; Barsky et al., 2013; Bermúdez (Turner, 1992; Martínez Navarro and Palmqvist, 1996; Arribas and de Castro et al., 2013). Even a cursory review of the available evi- Jorda, 1999; Arribas and Palmqvist, 1999; O'Regan, 2008; Made dence suggests a “virtual” scarcity of human presence in the Early and Mateos, 2010; Palombo, 2010; Made et al., 2011; O'Regan Pleistocene. According to this argumentation, hominin occupation et al., 2011). Very recently new approaches and hypothesis con- of Europe was occasional, infrequent and intermittent during the cerning the palaeoecological scenario of the human settlement Calabrian (1.4e0.8 Ma). joined the general discussion. Factors like faunal turnover, carni- In summary, beneath the debate summarized above is the im- vore guild structure, dynamics of the Pleistocene food webs and plicit idea that the evidence supporting the human population of resource availability (Arribas and Palmqvist, 1999; Croitor and Europe during the Early Pleistocene is scarce. Moreover, many Brugal, 2010; Palombo, 2010, 2014; Rodríguez et al., 2012, 2013; scholars implicitly assume that, although present, humans were Rodríguez-Gomez et al., 2013; Rodríguez-Gomez et al., 2014) not a common nor abundant component of the European Fig. 1. Geographical distribution of sites with evidence of hominin presence (top) and local faunas from the time interval 1.4e0.8 Ma (bottom). Detailed information on the sites is provided in Tables 1 and 2. Black dots: reliable Early Pleistocene lithic assemblages or sites with hominin fossils, white dots “dubious” Early Pleistocene lithic assemblages; white squares: sites providing local faunas (see text). J. Rodríguez et al. / Quaternary International 389 (2015) 119e130 121 ecosystems. However, scarcity is a relative concept and, thus, the 2013), only LFs confidently dated to the interval 1.4 Mae0.8 Ma key question is whether Homo sp. was a common species, or not, in (Calabrian, Early Pleistocene) were included in the database. LFs comparison with the other large mammals inhabiting Europe at the were selected according to bio-stratigraphic correlations and nu- end of the Early Pleistocene. merical ages provided by the original sources, although numerical Commonness or rarity of an extinct species, Homo in particular, ages were given priority over bio-stratigraphic correlations. should be evaluated in the context of the palaeontological record. It is A total of 51 LFs from 34 sites met the criteria to be included in universally assumed that the lack of completeness and the spatial and the database (Fig. 1 and Table 1). The inclusion of Huescar 1 in this temporal heterogeneity of the fossil record may make a species tableshouldbebriefly commented on. Luminescence chronology appear to be rarer than it really was. This effect is especially important suggest a Middle Pleistocene age, around 400e500 Kyr, for the when different periods or geographical areas are compared. Here we sediments of the fossil bearing layers (Demuro et al., 2014), present a quantitative measure of the rarity of Homo in the European however it is included here because the fauna strongly suggests a Early Pleistocene communities relative to other species. Moreover, late Early Pleistocene age. It has been repeatedly pointed out that since rarity of a species at continental scale is influenced by the size of taxonomic inconsistency is a potential drawback in this kind of its distribution range, we also compare the size of the area of distri- compiled lists (Palombo, 2010; Rodríguez et al., 2013). To address bution of Homo in Europe during the late Early Pleistocene with the this problem, we reviewed all faunal lists and applied uniform area of the other large mammal species of this period. taxonomic criteria to obtain a taxonomically consistent database. We based our review on systematic revisions of groups by other 2. Material and methods authors (see Rodríguez et al., 2012 and references therein). In case of doubt about the validity of a species we adopted a conservative, A database including georeferenced late Early Pleistocene Local non-splitting criterion, as explained in Rodríguez et al. (2013).As Faunas (LFs) and sites with evidence of human presence