How rare was human presence in Europe during the Early Pleistocene?

How rare was human presence in Europe during the Early Pleistocene?

Jesús Rodríguez ^a

, Ana Mateos ^a , Jesús Angel Martín-Gonz alez ^b

¹ , Guillermo Rodríguez-G omez ^a

aCENIEH, Paseo Sierra de Atapuerca, 3, 09002 Burgos, Spain

bDpt. Matematicas y Computacion, Universidad de Burgos, Burgos, Spain

a r t i c l e i n f o

Article history:

Available online 30 December 2014 Keywords:

Rarity Hominin Early settlement Geographical range Occupancy

a b s t r a c t

Beneath the hot debate about the tempo and mode of thefirst human colonization of Europe is the 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- ropean ecosystems. Here we present a quantitative assessment of the rarity and commonness of the European large mammal species during the 1.4e0.8 Ma period, including hominins. Considering the palaeontological record only,Homowas not one of the most common species in Europe, but it may not be considered a rare species. In contrast, taking into consideration the archaeological record, hominins 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.

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1. Introduction

Timing and nature of the early human colonisation of Europe has been a hotly debated topic for a long time. Nowadays, there is a general consensus in assuming that humans dispersed to Western Europe during the late Early Pleistocene (Carbonell et al., 2010; Made and Mateos, 2010; Moncel, 2010; Palombo, 2010, 2013; Allu e et al., 2013). It is generally accepted that humans occupied Europe as early as 1.4 Ma as evidenced by the fossils and lithic industry recovered at Barranco Le on D (Spain) and Pirro Nord (Italy) (Arzarello et al., 2007; but see Muttoni et al., 2013, 2011; but see Rolland, 2013; Toro-Moyano et al., 2013). Beyond the debate about the phylogenetic relationships between those hominin European populations (Stringer, 2012; G omez-Robles, 2013; Lordkipanidze et al., 2013; Arsuaga et al., 2014), a new debate aroused in the last years concerning the continuity or discontinuity of this initial settlement (Dennell, 2003; Leroy et al., 2011; Muttoni et al., 2011; Bermúdez de Castro et al., 2013;

Mosquera et al., 2013; Martínez et al., 2014). Some theoretical evolutionary scenarios have been proposed including argumen- tations in pro of a continuous settlement (Bermúdez de Castro and Martin on-Torres, 2013; Garcia et al., 2013; Martínez et al., 2013) and/or supporting a discontinuous and intermittent occupation by populations constrained by climatic and ecological

uctuations (Dennell, 2003; Agustí et al., 2009; Dennell et al., 2011; Leroy et al., 2011; MacDonald et al., 2012). These debates re

ected the usual problems derived from the low completeness of the hominin fossil record, which represents a fragmented, poorly-dated set of evi- dences. Several issues related to the ecological and environmental context of this dispersal event, or the technological skills of these populations are still under discussion (Turner, 1992; Arribas and Palmqvist, 1999; Mosquera et al., 2013). Moreover, recent discov- eries and critical evaluations of the chronological framework and dispersal events related to the

rst human colonization of Europe (Muttoni et al., 2011, 2013; Par es et al., 2013; Rolland, 2013) show that key questions about the tempo and mode of this colonization remain unsolved.

Several models have been proposed supporting the idea that the

rst human colonisers were restricted to the Mediterranean area because they were unable to survive in mid-latitude Europe (Roebroeks and Kolfschoten, 1992; Dennell and Roebroeks, 1996;

Roebroeks, 2001; Dennell et al., 2010). Although many of the

*Corresponding author.

E-mail addresses:[email protected](J. Rodríguez),ana.mateos@cenieh.

es (A. Mateos), [email protected] (J.A. Martín-Gonzalez), guillermo.

[email protected](G. Rodríguez-Gomez).

1 Temporarily assigned to CENIEH.

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Quaternary International

j o u r n a l h o m e p a g e :w w w . e l s e v i e r . c o m / l o c a t e / q u a i n t

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evolutionary scenarios proposed take into account only the palaeoanthropological and/or archaeological evidence (Carbonell et al., 1999; Dennell et al., 2010, 2011; MacDonald et al., 2012;

Barsky et al., 2013; Bermúdez de Castro and Martin on-Torres, 2013; Mosquera et al., 2013), it becomes increasingly evident that some complex relationships existed between palaeoclimatic and palaeoecological changes and human dispersals and settling (Turner, 1992; Martínez Navarro and Palmqvist, 1996; Arribas and Jord a, 1999; Arribas and Palmqvist, 1999; O'Regan, 2008; Made and Mateos, 2010; Palombo, 2010; Made et al., 2011; O'Regan et al., 2011). Very recently new approaches and hypothesis con- cerning the palaeoecological scenario of the human settlement joined the general discussion. Factors like faunal turnover, carni- vore guild structure, dynamics of the Pleistocene food webs and resource availability (Arribas and Palmqvist, 1999; Croitor and Brugal, 2010; Palombo, 2010, 2014; Rodríguez et al., 2012, 2013;

Rodríguez-G omez et al., 2013; Rodríguez-G omez et al., 2014)

highly constrained opportunities of the settlement for early hominins.

All things considered, human occupation during the Early Pleistocene is currently represented in Europe by a short list of sites.

Many authors assume that the fragmentary archaeological and fossil records indicate only sporadic hominin presence or marginal settlement (MacDonald et al., 2012; Barsky et al., 2013; Bermúdez de Castro et al., 2013). Even a cursory review of the available evi- dence suggests a

virtual

scarcity of human presence in the Early Pleistocene. According to this argumentation, hominin occupation of Europe was occasional, infrequent and intermittent during the Calabrian (1.4

0.8 Ma).

In summary, beneath the debate summarized above is the im- plicit idea that the evidence supporting the human population of Europe during the Early Pleistocene is scarce. Moreover, many scholars implicitly assume that, although present, humans were 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 inTables 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).

ecosystems. However, scarcity is a relative concept and, thus, the key question is whether

sp. was a common species, or not, in comparison with the other large mammals inhabiting Europe at the end of the Early Pleistocene.

Commonness or rarity of an extinct species,

in particular, should be evaluated in the context of the palaeontological record. It is universally assumed that the lack of completeness and the spatial and temporal heterogeneity of the fossil record may make a species appear to be rarer than it really was. This effect is especially important when different periods or geographical areas are compared. Here we present a quantitative measure of the rarity of

in the European Early Pleistocene communities relative to other species. Moreover, since rarity of a species at continental scale is in

uenced by the size of its distribution range, we also compare the size of the area of distri- bution of

in Europe during the late Early Pleistocene with the area of the other large mammal species of this period.

2. Material and methods

A database including georeferenced late Early Pleistocene Local Faunas (LFs) and sites with evidence of human presence was compiled from published sources. Since our focus is on large mam- mals, only species in the orders Artiodactyla, Perissodactyla, Carnivora, Primates and Proboscidea were included. Mustelids were not included in the analyses because of their small size. The geographical extent of this database was restricted to longitude be- tween 10

W and 35

E and latitude between 36

N and 55

N (Fig. 1).

This extent excludes the high latitude areas of Europe, which are known to have been colonized only during the Late Pleistocene (Pavlov et al., 2001). Although a small part of Anatolia is included inside the extent of our database, data from this Asian peninsula were not collected. Since the oldest evidence of human presence in Europe dates to 1.4 Ma (Arzarello et al., 2007; Toro-Moyano et al.,

2013), only LFs con

dently dated to the interval 1.4 Ma

0.8 Ma (Calabrian, Early Pleistocene) were included in the database. LFs were selected according to bio-stratigraphic correlations and nu- merical ages provided by the original sources, although numerical ages were given priority over bio-stratigraphic correlations.

A total of 51 LFs from 34 sites met the criteria to be included in the database (Fig. 1 and Table 1). The inclusion of Huescar 1 in this table should be brie

y commented on. Luminescence chronology suggest a Middle Pleistocene age, around 400

500 Kyr, for the sediments of the fossil bearing layers (Demuro et al., 2014), however it is included here because the fauna strongly suggests a late Early Pleistocene age. It has been repeatedly pointed out that taxonomic inconsistency is a potential drawback in this kind of compiled lists (Palombo, 2010; Rodríguez et al., 2013). To address this problem, we reviewed all faunal lists and applied uniform taxonomic criteria to obtain a taxonomically consistent database.

We based our review on systematic revisions of groups by other authors (see Rodríguez et al., 2012 and references therein). In case of doubt about the validity of a species we adopted a conservative, non-splitting criterion, as explained in Rodríguez et al. (2013). As an effect of this decision our results may overestimate the oc- currences of some species. Several LFs included taxa identi

ed only at the genus level. We treated these cases as follows. If more than one species of the genus is recognized in this time period records at genus level were not included in the analyses. Other- wise, if a single species of the genus is recognized for this period, records identi

ed at the genus level are assimilated to the species.

As an example, all records reported as

sp. are included in the analyses as

sp./Macaca sylvanus. Finally, some taxa were only recorded at the genus level in the sites included in the database, e.g.

sp. These cases were treated as additional species. Again, these criteria tend to overestimate the occurrences of those taxa affected.

Table 1

Late Early Pleistocene (1.4e0.8 Ma) local faunas included in the analyses.

Locality Longitude Latitude Age Reference

1 Sima del Elefante TE 09 3.52 42.35 1.4e1.2 Ma (Rodríguez et al., 2011)

2 Barranco Leon D 2.43 37.71 1.4e1 Ma (Toro-Moyano et al., 2013)

3 Cueva Victoria 1.18 37.85 1.4e0.9 Ma (Gibert, 1993; Gibert et al., 1999)

4 Pirro Nord 13.65 43.28 1.4e1.2 Ma (Palombo et al., 2000e2002; Arzarello et al., 2007)

5 Sandalja-I 13.89 44.90 1.4e1.2 Ma (Kahlke et al., 2011)

6 Ceyssaguet 1 3.90 45.12 1.3e1.2 Ma Paleobiology Database Searched 16-04-2010

7 Saint-Prest 1.50 48.36 1.3e0.8 Ma (Guerin et al., 2003)

8 Sierra de Quibas 0.93 38.20 1.3e1.07 Ma (Montoya et al., 1999; Montoya et al., 2001)

9 Sima del Elefante TE 10 3.52 42.35 1.2e1 Ma (Rodríguez et al., 2011)

10 Sima del Elefante TE 11 3.52 42.35 1.2e1 Ma (Rodríguez et al., 2011)

11 Sima del Elefante TE 12 3.52 42.35 1.2e1 Ma (Rodríguez et al., 2011)

12 Sima del Elefante TE 13 3.52 42.35 1.2e1 Ma (Rodríguez et al., 2011)

13 Sima del Elefante TE 14 3.52 42.35 1.2e1 Ma (Rodríguez et al., 2011)

14 Apollonia-1 23.44 40.62 1.2e0.8 Ma (Kostopoulus, 1997)

15 Betfia level 3b 22.03 46.97 1.2e0.8 Ma Paleobiology Database Searched 16-04-2010

16 Durfort 3.95 43.98 1.2e1 Ma (Palombo and Valli, 2003e2004)

17 Huescar-1 2.50 37.77 1.2e0.8 Ma (Diago et al., 2003)

18 Imola 11.71 44.35 1.2e0.99 Ma (Palombo et al., 2000e2002)

19 Kozarnika 11c 22.68 43.63 1.2e0.8 Ma (Guadelli et al., 2005)

20 Kozarnika 11d 22.68 43.63 1.2e0.8 Ma (Guadelli et al., 2005)

21 Kozarnika 12 22.68 43.63 1.2e0.9 Ma (Guadelli et al., 2005)

22 Kozarnika 13 22.68 43.63 1.2e0.78 Ma (Guadelli et al., 2005)

23 Kunino Lower Level 23.96 43.18 1.2e0.99 Ma (Palombo and Mussi, 2006)

24 Marathousa-Megalopolis 22.04 37.51 1.2e0.99 Ma (Palombo and Mussi, 2006)

25 Ravin Voulgarakis 23.50 40.60 1.2e0.99 Ma (Palombo and Mussi, 2006)

26 Rosieres“Usine” 2.25 46.95 1.2e0.99 Ma (Despriee et al., 2007)

27 Torre di Picchio 12.49 42.65 1.2e1 Ma (Raia et al., 2009)

28 Redicicoli 12.52 41.93 1.2e0.8 Ma (Raia et al., 2009)

29 Dealul Viilor 24.66 45.15 1.1e0.9 Ma (Palombo and Mussi, 2006)

30 Sainzelles 3.80 45.08 1.1e0.99 Ma (Palombo and Valli, 2003e2004)

31 Colle Curti 12.75 42.95 1.07e0.78 Ma (Palombo et al., 2000e2002)

32 Grotte du Vallonnet BI 7.47 43.77 1.07e1 Ma (Lumley et al., 1988b)

(continued on next page)

Evidence of human presence in Europe during the Early Pleis- tocene both in the form of fossils and lithic artifacts was compiled from published sources. Several of those sites allegedly evidencing an old human presence in Europe are controversial, thus some of them were

agged as

dubious

on the basis of the criticisms raised by other authors (Table 2).

Frequency of occurrence in the palaeontological record, or prevalence, a measure of occupancy (Gaston and Fangliang, 2010),

is taken as a proxy for the commonness of a species during the Early Pleistocene inside the sampled area. The prevalence (P) of a species is computed as the number of LFs that contain this species.

We also compared the amplitude of the geographical distribu- tions of

and other Early Pleistocene large mammals. Longi- tudinal amplitude (LgA) was computed as the maximum difference

in longitude between two sites where the species is present.

Similarly, latitudinal amplitude (LtA) is de

ned as the maximum

Locality Longitude Latitude Age Reference

33 Grotte du Vallonnet BII 7.47 43.77 1.07e1 Ma (Lumley et al., 1988b)

34 Grotte du Vallonnet C 7.47 43.77 1.07e1 Ma (Lumley et al., 1988b)

35 Grotte du Vallonnet III 7.47 43.77 1.07e1 Ma (Lumley et al., 1988b)

36 K€arlich Level A 7.47 50.47 1.07e0.9 Ma Paleobiology Database Searched 16-04-2010

37 Soleilhac 4.02 45.02 1.07e0.9 Ma (Alberdi et al., 1997; Palombo and Valli, 2003e2004)

38 Untermassfeld 10.42 50.55 ~1.07 Ma (Koenigswald and Heinrich, 1999)

39 Gran Dolina TD5 3.52 42.35 1e0.8 Ma (Rodríguez et al., 2011)

40 Betfia level 3c 22.03 46.97 1e0.8 Ma Paleobiology Database Searched 16-04-2010

41 Chlum 6 16.76 49.41 1e0.9 Ma (Koenigswald and Heinrich, 1999)

42 Incarcal 1 2.78 42.19 1.4e0.9 Ma (Julia, 1984; Galobart, 2007; Ros-Montoya et al., 2012)

43 Het Gat 3.22 52.40 1e1.2 Ma (Mol et al., 2003)

44 Gran Dolina TD3eTD4 3.52 42.35 0.99e1 Ma (Rodríguez et al., 2011)

45 Estrecho de Quipar 1.88 38.04 0.99e0.9 Ma (Scott and Gibert, 2009)

46 K€arlich Level B 7.47 50.47 0~0.99 Ma Paleobiology Database Searched 16-04-2011

47 Vallparadís 10-10c 2.02 41.56 0.98e0.78 Ma (Martínez et al., 2010; Madurell-Malapeira et al., 2011)

48 Gran Dolina TD6 1-2 3.52 42.35 ~0.9 Ma (Rodríguez et al., 2011)

49 Gran Dolina TD6-3 3.52 42.35 ~0.9 Ma (Rodríguez et al., 2011)

50 Happisburgh Site 3 D-E 1.53 52.82 ~0.8 Ma (Parfitt et al., 2010)

51 Fuente Nueva 3 2.4 37.71 1.19±0.21 (Martínez-Navarro et al., 2010)

Table 2

Palaeontological and lithic assemblages providing evidence of the presence of hominins in Europe earlier than 0.8 Ma. Code for dating methods as follows: BS: biostratigraphic correlation, ESR: electron spin resonance, ESR-OB: electron spin resonance of optically bleached quartz grains, CN: cosmogenic nuclides, PM: paleomagnetism, TL:

Thermoluminescence.

Site Long. Lat. Technology Hominin

fossils

Estimated age (Ma)

Dating methods

Reference

Reliable evidence

Kozarnika (Bulgaria) 22.68 43.63 Mode 1 1.4e1.6 BS (Sirakov et al., 2010)

Pirro Nord P13 (Italy) 13.65 43.28 Mode 1 1.3e1.7 BS (Arzarello et al., 2007)

Barranco Leon D (Spain) 2.43 37.71 Mode 1 Homosp. 1.43±0.38 BS, PM, ESR (Toro-Moyano et al., 2009;

Toro-Moyano et al., 2013)

Atapuerca Sima del Elefante TE9 (Spain) 3.52 42.35 Mode 1 Homosp. 1.4e1.2 CN (Pares et al., 2006; Carbonell et al., 2008) Atapuerca Sima del Elefante TE11 (Spain) 3.52 42.35 Mode 1 1.2e0.78 CN, PM (Rodríguez et al., 2011; Olle et al., 2013) Atapuerca Sima del Elefante TE12 (Spain) 3.52 42.35 Mode 1 1.2e0.78 CN, PM (Rodríguez et al., 2011; Olle et al., 2013) Atapuerca Sima del Elefante TE13 (Spain) 3.52 42.35 Mode 1 1.2e0.78 CN, PM (Rodríguez et al., 2011; Olle et al., 2013) Atapuerca Sima del Elefante TE14 (Spain) 3.52 42.35 Mode 1 1.2e0.78 CN, PM (Rodríguez et al., 2011; Olle et al., 2013) Fuente Nueva 3 (Spain) 2.4 37.71 Mode 1 1.19±0.21 PM, BS, ESR (Oms et al., 2000; Toro-Moyano et al., 2009;

Duval et al., 2012)

Vallparadís EVT8-7 (Spain) 2.02 41.56 Mode 1 0.98e0.95 PM, BS, ESR (Martínez et al., 2010; Duval and Moreno, 2011;

Madurell-Malapeira et al., 2012;

Martínez et al., 2014)

Atapuerca Gran Dolina TD6 (Spain) 3.52 42.35 Mode 1 H. antecessor 0.9 PM, BS, ESR (Rodríguez et al., 2011; Pares et al., 2013) Atapuerca Gran Dolina TD3eTD4 (Spain) 3.52 42.35 Mode 1 0.94±0.1 ESR-OB,

BS, TL

(Moreno, 2011; Rodríguez et al., 2011)

Pont de Lavaud (France) 1.88 46.25 Mode 1 1.07±0.09 ESR (Bahain et al., 2007)

Terre des Sablons

(Lune Rosiers Unit 3, France)

2.26 46.95 Mode 1 1.1±0.18

to 930

ESR (Despriee et al., 2010)

Le Vallonet (France) 7.46 43.77 Mode 1 >0.9z1.0 ESR, PM, BS (Yokoyama et al., 1988; Lumley et al., 1988a;

Gagnepain, 1996)

Pont de la Hulauderie (France) 1.2 47.92 Mode 1 0.98 ESR (Despriee et al., 2010)

Happisburgh (UK) 1.53 52.82 Mode 1 Footprints 0.98e0.78 PM, BS (Parfitt et al., 2010; Ashton et al., 2014)

Korolevo VII (Ukraine) 23.13 48.15 Mode 1 0.95 PM (Koulakovska et al., 2010)

Monte Poggiolo (Italy) 12.04 44.22 Mode 1 1.06±0.16 ESR (Bahain et al., 2007)

“Dubious”evidence

Lezignan Le-Cebe (France) 3.43 43.48 Mode 1 1.57 ³⁹Ar/⁴⁰Ar, BS (Crochet et al., 2009)

Anagni (Italy) 13 41.77 Mode 1 >0.706 KeAr (Biddittu et al., 1979)

Colle Marino-Arce-Fontana Liri (Italy) 13.54 41.62 Mode 1 >0.706 e (Biddittu et al., 1979)

Untermassfeld (Germany) 10.42 50.55 Mode 1 1.05 BS, PM (Landeck, 2010; Garcia et al., 2013)

Dorn-Dürkheim 3 (Germany) 8.26 49.76 Mode 1 >0.78 PM (Haidle and Pawlik, 2010)

K€arlich A (Germany) 7.49 50.39 Mode 1 1.07e0.99 PM (Bosinski et al., 1980; Würges, 1986)

Estrecho del Quípar (Spain) 1.88 38.04 Mode 2 c.0.9 PM (Scott and Gibert, 2009)

difference in latitude between two sites where the species is pre- sent. Both variables LgA and LtA are expressed in degrees. Range size (Rg) is used as a proxy for the geographical range size of the species. It is computed as the maximum distance in km between two sites where the species is present. The distance between each pair of sites was computed based on their latitude and longitude.

The geodesic distance, i.e. the length of the shorter arc of the great circle passing through both sites, was computed assuming the Earth is a sphere, instead of using a spheroid to account for the

attering of the Earth.

Carnivore and primary consumers are known to have different allometric constraints in several parameters related to their dis- tribution, like population density (Damuth, 1981) or home range size (Martin, 1991). Thus, carnivore and primary consumer species were analyzed separately. Moreover, since the three LFs which

contain human fossils are located in the Iberian Peninsula we car- ried out our analyses at two different extents: the Iberian Peninsula and the entire sampled area. Although it excludes high latitudes and the land to the east of 35

E longitude, from here on out we will refer to the later area as Europe.

Empirical Cumulative Density Functions (CDF) of P have been computed and used to calculate in which quartile of the distribu- tion the prevalence of

is included. In ecology, species with an abundance or distribution range below the median are considered to be rare (Yu and Dobson, 2000). Here the quartile is taken as an index of the rarity or commonness of

and other species in the Early Pleistocene. Species in the

rst quartile are considered rare, those in the second quartile are considered moderately rare, the third quartile includes the moderately common species and the fourth quartile the common species. A Kolmogorov

Smirnov test

Fig. 2.Prevalence of carnivores (top) and other large mammal species (bottom) in Europe during the late Early Pleistocene. Hominin prevalence estimated from the number of sites with fossils attributed to the genusHomoand from the number of lithic assemblages is compared with both distributions. The white part in the bar of lithic assemblages represents dubious assemblages (see text).

was used to test for differences in the distribution of geographical range size between carnivores and other large mammals. Statistical analyses were carried out using Matlab R2009b.

3. Results

3.1. Evidence of human presence in the late Early Pleistocene

Fossils attributed to the genus

have been found only in three Early Pleistocene sites: Sima del Elefante TE9, Gran Dolina TD6 and Barranco Le on D, all of them located in the Iberian Peninsula. Though, this restricted distribution of human fossils contrasts with the wide distribution of the lithic record from the same period (Table 2). A total of 26 European lithic assemblages have been attributed an age older than 0.8 Ma, although 7 of them are considered dubious. These later localities contain lithic and faunal assemblages in an ambiguous status, either imprecise

chronologies, geological and stratigraphic contexts or collections of uncertain archaeological nature. The precise stratigraphic prove- nance of the lithic artifacts from L ezignan Le-C ebe (France) (Crochet et al., 2009) is problematic (Muttoni et al., 2011; Par es et al., 2013; Rolland, 2013). The artifacts of the Italian localities of Anagni and those of the Latium area (Colle Marino, Arce, Fontana Liri) (Biddittu et al., 1979) have an imprecise archaeological context and have been dated by regional geological correlations (Rolland, 2013). The German sites of Untermassfeld (Landeck, 2010; Garcia et al., 2013), Dorn-Dürkheim 3 and K€ arlich A (Haidle and Pawlik, 2010) are also problematic. The lithic artifacts from Untermass- feld were not found in association with the well-known fauna from this palaeontological site (Landeck, 2010). Likewise, the strati- graphic context of the layer K€ arlich A is also doubtful (Roebroeks and van Kolfschoten, 1995; Baales et al., 2000). The nature of the artifacts recovered at Dorn-Dürkheim 3 and K€ arlich A is contro- versial (Baales et al., 2000; Haidle and Pawlik, 2010 and references

Fig. 3.Prevalence of carnivores (top) and other large mammal species (bottom) in the Iberian Peninsula during the late Early Pleistocene. Hominin prevalence estimated from the number of sites with fossils attributed to the genusHomoand from the number of lithic assemblages is compared with both distributions. The white part in the bar of lithic assemblages represents dubious assemblages (see text).

therein; Roebroeks and van Kolfschoten, 1995). In Spain, the mag- netostratigraphy and the correlations based on the fauna of Cueva Negra del Estrecho del Quípar (Scott and Gibert, 2009) remain questionable (Jim enez-Arenas et al., 2011; Muttoni et al., 2011).

3.2. Prevalence of Homo in the context of the palaeontological record

A total of 83 mammal species, 29 of them carnivores, were included in the database. Prevalence at continental scale ranges from 1 to 18 occurrences for the order Carnivora and from 1 to 19 occurrences for other mammalian orders (Fig. 2). In contrast, P ranges from 1 to 10 for carnivores and from 1 to 12 for other mammals when only the LFs from the Iberian Peninsula are considered (Fig. 3). The distribution of P is severely skewed to the left, as shown in Fig. 4, irrespectively of the taxonomic group or the geographical extent.

Taking into account only the palaeontological record,

shows a low prevalence (3 sites). Though considering the empiric distribution of P (Fig. 4c), the prevalence of

is included in the third quartile of the prevalence of primary consumers in Europe and in the second quartile of the prevalence of carnivores.

Considering only the Iberian Peninsula the prevalence of

is also in the third quartile of the prevalence of primary consumers and in the second quartile of carnivores (Fig. 4f).

In contrast, if the lithic record is taken into consideration, humans become one of the most common species in Europe with 15 reliable and 7 dubious occurrences (Fig. 2). In the Iberian Peninsula the lithic record also places humans among the most common species with 6 reliable and one dubious record (Fig. 3).

Considering the reliable lithic records only, the prevalence of humans would be on the fourth quartile of the distributions of

primary consumers and carnivores both in Europe and in the Ibe- rian Peninsula (Fig. 4c and f).

3.3. Range size

The distribution of the range sizes of the late Early Pleistocene large mammal species is severely skewed to the left (Fig. 5). This is a direct consequence of the distribution of P, since the range of the species occurring in a single site is 0. At

rst glance, carnivores seem to exhibit a bimodal distribution of Rg with a tendency to- wards larger range size in the species recorded in more than one site. In contrast, the distribution of range sizes seems to be more homogeneous in primary consumers (Fig. 5). However, this apparent differences in range size between carnivores and primary consumers are not statistically signi

cant (K

S test

0.3048,

0.06).

Fig. 6 shows the longitudinal and latitudinal amplitudes of the late Early Pleistocene large mammals in Europe. Again, a tendency towards wider geographical ranges is evident for carnivores.

Because all hominin fossils from this period have been found in the Iberian Peninsula, the latitudinal and longitudinal amplitude of

is very small when computed in this way. However, the amplitude of the distribution of lithic assemblages is one of the largest for any large mammal in the sample (Fig. 6). The latitudinal and longitudinal amplitude of the lithic record has been obtained considering reliable assemblages only.

4. Discussion

Commonness and rarity are complex concepts in Ecology that involve local population density, trophic level, geographical dis- tribution range, and variety of habitats occupied by a species

Fig. 4.Distribution of the prevalence of primary consumers (a), (d) and carnivores (b), (e) in Europe (upper row) and the Iberian Peninsula (lower row). The cumulative density functions (CDFs) of the distributions of prevalence in Europe (c) and the Iberian Peninsula (f) are used to evaluate the commonness or rarity of hominins in the late Early Pleistocene (see text). Continuous line: primary consumers; dashed line: carnivores. The vertical dotted lines show the prevalence of hominins computed on the basis of the fossil record and on the basis of the number of reliable lithic assemblages.

(Rabinowitz, 1981; Gaston and Fuller, 2007; Bennett and Provan, 2008; Gaston, 2008). Population density is dif

cult to estimate on the basis of palaeontological data, but see Meloro et al. (2007).

Thus, we focus our evaluation of rarity in late Early Pleistocene large mammals on the number of occurrences (prevalence) and the size of the distribution range. The pattern of rarity observed in our results, either for carnivores or primary consumers, is composed of a small number of common species and a large number of rare species. This pattern is not an artefact of the palaeontological re- cord, as may be thought, since the same pattern is observed in recent faunas (Gaston, 2008). In her classic book on biological di- versity Magurran (2004, pp. 18) wrote:

In no environment

are all species equally common. Instead, it is universally the case that some are abundant, others only moderately common, and the remainder-often the majority-rare

. Rarity is not unusual among recent large mammals. Almost half the species of carnivores in the world have relatively small populations and distributions, and mammals as a whole show a high frequency of rare species (Yu and Dobson, 2000).

Although the distribution of prevalence shown by our results is coherent with what may be expected on the basis of the frequency of rarity among recent mammals, it should be acknowledged that our results may have been affected by the faunal turnover occurred inside the time period under consideration. Most of the 1.4

0.8 Ma time interval corresponds to the early Galerian or Epi- Villafranchian Large-Mammal Age (Palombo, 2010; Kahlke et al., 2011). Nevertheless, the older LFs included in our database should be considered as late Villafranchian (like Pirro Nord, Sandalja-1, Ceyssaguet 1, Sainzelles, Beftia or Barranco Leon D).

Consequently, some of the species included in the analyses were not present in Europe along the entire time interval considered.

Likely, some of those species were widely distributed and common, but because of their restricted time distribution they are recorded only in the local faunas of appropriate age. Those species would seem to be rarer than they actually were. In despite of this possible bias, restriction of the analyses to the Epi-Villafranchian faunas has little effect on the pattern of prevalence and on the distribution of geographical range sizes (results not shown).

On average, late Early Pleistocene carnivores exhibit higher prevalence than primary consumers (Fig. 4). This result may seem to be odd, because it is a well-established rule that carnivores have lower population densities than primary consumers of the same size (Damuth, 1991; Carbone and Gittleman, 2002). However, prevalence is not related to population density or abundance in a simple way. A species may be common because it has a large dis- tribution and it is found in many localities inside its distribution, although at low densities. In his analysis of rarity in recent mam- mals Yu and Dobson (Yu and Dobson, 2000) found that almost 25%

of the mammal species with broad habitat distributions occur at low population densities. The relatively high prevalence of late Early Pleistocene carnivores is likely related to the larger sizes of their geographical ranges. It has been widely demonstrated that carnivores have lower population density and larger home ranges than primary consumers of similar size (Kelt and Van Vuren, 1999;

Jetz et al., 2004). Thus, carnivores must have large geographical range sizes to sustain populations large enough to avoid extinction (Hern andez Fern andez and Vrba, 2005). Moreover, the relatively large geographical range of carnivores is also related to their increased home ranges, marked territoriality and high dispersal abilities. Our results failed to detect statistically signi

cant differ- ences in the sizes of the geographical ranges of carnivores and primary consumers, but the distribution of Rg (Fig. 5) and of the latitudinal and longitudinal amplitude (Fig. 6) suggest that a ten- dency exists in carnivores towards larger geographical ranges.

4.1. Commonness vs. rarity in hominins

It is apparent from our results that, if the rarity of humans in late Early Pleistocene Europe is judged on the basis of the number of sites with hominin fossils,

was not one of the most common taxa in the European ecosystems.

appears as a moderately rare taxon when compared with the prevalence of carnivores, but not when compared with the prevalence of primary consumers. In contrast, in the Iberian Peninsula, where all sites with

re- mains are located, the prevalence of humans is above the median of either carnivores or primary consumers and, thus, it may be considered a moderately common species. Although the trophic behaviour of Early Pleistocene hominins is a hotly debated topic (Binford, 1981; Blumenschine and Madrigal, 1993; Bunn and Ezzo, 1993; Bunn, 2001; Pickering, 2001; Bunn and Pickering, 2010;

Domínguez-Rodrigo et al., 2010; Bunn and Gurtov, 2014), they are best described as omnivores that included a high proportion of animal food in their diets, and used large mammals as a key trophic resource Indeed, it has been suggested that hominins were skilled scavengers, able to successfully compete with such a highly specialized carnivore as the giant hyaena (Espigares et al., 2013) Thus, hominins are expected to behave in an intermediate way between strict primary and secondary consumers. Taking all this together it is apparent that, without being an extremely common taxon,

sp. was not a rare component of the late Early Pleis- tocene ecological communities of Europe.

The question of how many hominin species inhabited Europe during the Early Pleistocene is beyond the scope of this paper.

Though, it is necessary to acknowledge that existence of more than one hominin species in Europe during this period, as proposed by some authors (Bermúdez de Castro and Martin on-Torres, 2013),

Fig. 5.Distribution of the range size (Rg) in km of large primary consumers (a) and carnivores (b). The empirical cumulative density functions (c) have been used to test for differences between the distributions using a KolmogoroveSmirnov test.

would affect our results. If the European Early Pleistocene

fossils actually correspond to several species the prevalence of each of those hominin species would be markedly lower than estimated for the genus. Moreover, it would not be possible to match a lithic assemblage with a particular species, and our analyses of the size of the distribution range would not be valid. Analyses of rarity in Early Pleistocene mammals at the genus level may be proposed as an alternative to address this eventuality. However, an analysis at the genus level would have little ecological meaning, since rare and common species in the same genus would be merged together. In addition, since not all genera include the same number of species, those genera with more species would tend to exhibit higher prevalence. Moreover, as explained in the material and methods section, we adopted a conservative, non-splitting, criterion in reviewing the taxonomic consistency of the large mammal LFs.

Thus, there is no reason to change the criterion in the case of hominins. Taking into account all these considerations, and given than it is currently not known how many hominin species inhabi- ted Europe, we present our results at the species level as a

rst quantitative evaluation of the relative rarity of humans in the late Early Pleistocene.

When their prevalence and distribution in late Early Pleistocene Europe is estimated from the lithic archaeological record, hominins become a common and widely distributed species. However, the lithic and the palaeontological records are of different nature, and a direct comparison is not warranted. In general, lithic artifacts have higher preservational probabilities than bones. Moreover, a single hominin individual may contribute to the palaeontological record not only with a single skeleton, composed of several elements, but also with thousands of lithic artefacts produced during its life. Thus, we must expect lithic evidence always to be more abundant than palaeontological evidence. In any case, the lithic archaeological record shows that hominins were one of the most widely distrib- uted European species in the late Early Pleistocene. More inter- estingly, hominins were one of the few species with a wide latitudinal range, may be an indication of a highly carnivorous diet.

Certainly, variations in the distribution of species in Europe during the late Early Pleistocene cannot be assessed with the data pre- sented here. Thus, the geographical ranges estimated here should be taken as the maximum distribution of the species in the period as a whole. Likely, many species never occupied simultaneously the

entire range estimated for them, but this was not the case of hominins. The lithic record show that hominins were present at Happisburgh, Atapuerca and Korolevo VII (Koulakovska et al., 2010;

Par

tt et al., 2010; Rodríguez et al., 2011) at roughly the same time.

The lithic record suggests that humans had a high occupancy and were widely distributed across Europe during the late Early Pleistocene, but this does not imply that they were abundant. As discussed above, prevalence and geographical range size are not directly related to local population density. Moreover, a low pop- ulation density would explain why hominins show a moderate prevalence in the palaeontological record. We lack reliable proxies to estimate local population densities in the past. Number of ele- ments or minimum number of individuals may serve to assess the relative abundance in the environment of the species in a fossil assemblage (Delpech, 1999; Hertler and Volmer, 2008; Fa et al., 2013), although taphonomic processes may severely affect the es- timates. Comparison of species abundances among different as- semblages, affected by diverse accumulation and diagenetic processes, is even more problematic using those proxies. Recently, Rodríguez-G omez et al. (2013) explored resource availability at Atapuerca TD6 and showed that this southern European ecosystem was able to sustain high hominin population densities. Moreover, Rodríguez et al. (2014) showed that ungulate carrying capacity was high in Mediterranean ecosystems during the late Early Pleisto- cene, suggesting abundance of meat resources. Since all hominin fossils have been found in southern Europe, and assuming that the probability of being preserved in the fossil record is directly pro- portional to population density, it may be speculated that hominins occurred at higher population densities in southern Europe than in the rest of the continent. An analysis of the spatial variation of resource availability across Europe during the late Early Pleistocene and its relationship to the distribution of hominin occupancy would help to test this hypothesis.

5. Conclusions

The palaeoanthropological record suggests that hominins were not a rare component of the late Early Pleistocene European eco- systems, although they were not among the most common ele- ments. In contrast, the archaeological record suggests that they were widely distributed. It may be speculated that hominins were present in many areas but at low densities. On the one hand, if humans occurred at low population densities their probability to be preserved as fossils would be low. On the other hand, a small hu- man population would be able to produce several lithic assem- blages, some of which might be recovered. Considering Early Pleistocene hominins as a frequent but not abundant species adds a new dimension to our understanding of the ecological and de- mographic constraints of the

rst colonization of Europe.

Acknowledgements

This research was funded by the MINECO project, CGL2012- 38434-C03-02. G. Rodríguez-G omez was the bene

ciary of a pre- doctoral FPI Grant from the Spanish MICINN. We thank L. Arnold, M. Duval, D. Hoffmann and J. M. Par es for having invited J. Rodrí- guez to participate in the

The Early-Middle Pleistocene transition:

Signi

cance of the Jaramillo subchron in the sedimentary record

Workshop and to contribute to this volume. Thanks also to the workshop participants for their thought provoking discussion and suggestions and especially to B. Martínez-Navarro and an anony- mous reviewer for their useful comments on the manuscript. An anonymous editor from Elsevier's Language services improved the English of the original manuscript.

Fig. 6.Latitudinal and longitudinal amplitude of the geographical distribution of large primary consumers (squares), carnivores (black dots) and hominins (black stars) in Europe during the late Early Pleistocene. The geographical distribution of hominins has been calculated both on the basis of the fossil record (F) and on the basis of lithic assemblages (L). The dotted lines indicate the maximum possible latitudinal and longitudinal differences between two points inside the sampled area.

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