European Early Pleistocene Research Articles

Increase on environmental seasonality through the European Early Pleistocene inferred from dental enamel hypoplasia

An in-depth study of the Early Pleistocene European remains of Hippopotamus has allowed the first detailed description of the incidence and types of dental alterations related to palaeopathologies and potentially linked to climatic and environmental factors. The results of a long-term qualitative and quantitative assessment highlight the importance of nutrient deficiencies on the development of dental enamel hypoplasia in Hippopotamus. Glacial cyclicity and the resulting changes in humidity and plant community structure conditioned the local environments critical for the survival of this taxon. Two main intervals of putative constrained nutritionally restrictions were detected at ca. 1.8 Ma and ca. 0.86 Ma (i.e., MIS63 and MIS21, respectively). Statistical comparisons show an increase in the frequency of dental hypoplasia between these two chronological periods, thus reinforcing the idea of increased seasonality in the circum-Mediterranean environments during the Early Pleistocene.

The Technological Behaviours of Homo antecessor: Core Management and Reduction Intensity at Gran Dolina-TD6.2 (Atapuerca, Spain)

The ability of early hominins to overcome the constraints imposed by the characteristics of raw materials used for stone tool production is a key topic on the discussion about the evolution of hominin cognitive capabilities and technical behaviours. Thus, technological variability has been the centrepiece on this debate. However, the variability of lithic assemblages cannot be correctly interpreted without understanding site occupational models and function and considering that individual tools represent specific discard moments in a continuous reduction process. In Europe, the earliest technological record is represented by the scarce and scattered Mode 1 technologies, often deriving from occasional occupations or restricted activity areas yielding unrepresentative assemblages. In this paper, we approach the technological behaviours exhibited by Lower Palaeolithic hominins from the subunit TD6.2 of the Gran Dolina site (Atapuerca, Burgos) by including the perspective of reduction intensity studies on the analysis of technological variability. Gran Dolina TD6.2 is a unique and extremely significant archaeological context, as it represents the oldest multi-layered unit of domestic hominin occupations in the Early Pleistocene of Europe. We use the Volumetric Reconstruction Method (VRM) to estimate the original volume of the blanks and quantify the reduction intensity of each core individually to characterise the reduction distribution patterns using Weibull probability distribution functions. Our results suggest differential raw material management in terms of reduction intensity, according to the characteristics of each lithology. This could reflect a solid understanding of raw material qualities and a certain degree of planning. Altogether, the continuity between knapping strategies through reduction denotes constant adaptation to raw material constraints as well as particular knapping conditions, rather than specific compartmentalised mental schemes. In conclusion, Homo antecessor toolmakers would have been situational knappers whose technological behaviour would be highly adaptive. This research constitutes the first reduction approach for the European Early Pleistocene assemblages that will lead to a referential framework for other European Early Pleistocene sites.

<em>EQUUS STEHLINI</em> AZZAROLI, 1964 (PERISSODACTYLA, EQUIDAE). A REVISION OF THE MOST ENIGMATIC HORSE FROM THE EARLY PLEISTOCENE OF EUROPE, WITH NEW INSIGHTS ON THE EVOLUTIONARY HISTORY OF EUROPEAN MEDIUM- AND SMALL-SIZED HORSES

The present work provides a revision of the complete fossil collection of Equus stehlini from the Italian Early Pleistocene (late Villafranchian) of the Upper Valdarno Basin (Tuscany, Italy). The species was originally described in the second half of the last century, even if recent studies questioned its distinct species status, considering it a subspecies of Equus senezensis (i.e., E. senezensis stehlini). Over the succeeding decades the debate about the taxonomic status of E. stehlini extended to its possible evolutionary origin from two different Equus species from the European Early Pleistocene, Equus stenonis or E. senezensis. The revision undertaken here of E. stehlini, by means of morphological, morphometric and statistical comparisons, provides new insights into the taxonomic status and the evolutionary history of this enigmatic species. It is concluded that E. stehlini is a distinct species derived from the medium-sized E. senezensis. This work further highlights new insights and perspectives in the evolutionary framework and in the paleoecological understanding of the medium to small-sized Equus species from the early to middle Early Pleistocene of Europe.

An updated review on Equus stenonis (Mammalia, Perissodactyla): New implications for the European early Pleistocene Equus taxonomy and paleoecology, and remarks on the Old World Equus evolution

Equus stenonis is one of the most iconic fossil species of the Early Pleistocene. Although it is known from the 19th century, its taxonomic and evolutionary position has been extensively debated among equid specialists, including multiple interpretations. The current work aims to review the European E. stenonis record, by morphological, morphometrical and statistical analyses combined with body mass and net primary production estimates. Furthermore, we investigate the evolution of E. stenonis cranial anatomy in comparison with the Plio-Pleistocene species from North America, Eurasia and Africa, and extant zebras and asses. Our review of European E. stenonis samples has closely considered the E. stenonis subspecies Equus stenonis vireti, Equus stenonis guthi, Equus stenonis pueblensis, Equus stenonis olivolanus and Equus stenonis stenonis which we recognize as being ecomorphotypes of Equus stenonis, with the exception of Equus stenonis senezensis and Equus stenonis mygdoniensis. These two last subspecies warrant the species ranks of Equus senezensis and Equus altidens, respectively. This has led us to propose a new revised biochronology of European Early Pleistocene Equus. Our evolutionary interpretations based on cranial morphology reveals that the evolutionary lineage of extant zebras can be recognized in deep time from the North American species Equus simplicidens, through the European E. stenonis and the African Equus koobiforensis. European stenonine horses likewise represent the original source for the African Pleistocene fossil species and extant Equus grevyi.

Uneven Data Quality and the Earliest Occupation of Europe\u2014the Case of Untermassfeld (Germany)

The database regarding the earliest occupation of Europe has increased significantly in quantity and quality of data points over the last two decades, mainly through the addition of new sites as a result of long-term systematic excavations and large-scale prospections of Early and early Middle Pleistocene exposures. The site distribution pattern suggests an ephemeral presence of hominins in the south of Europe from around 1 million years ago onward, with occasional short northward expansions along the western coastal areas when temperate conditions permitted. From around 600,000–700,000 years ago, Acheulean artefacts appear in Europe and somewhat later hominin presence seems to pick up, with more sites and now some also present in colder climatic settings. It is again only later, around 350,000 years ago, that the first sites show up in more continental, central parts of Europe, east of the Rhine. A series of recent papers on the Early Pleistocene palaeontological site of Untermassfeld (Thuringia, Germany) makes claims that are of great interest for studies of earliest Europe and are at odds with the described pattern: the papers suggest that Untermassfeld has yielded stone tools and humanly modified faunal remains, evidence for a 1 million years old hominin presence in European continental mid-latitudes, and additional evidence that hominins were well-established in Europe already around that time period. Here, we evaluate these claims and demonstrate that these studies are severely flawed in terms of data on provenance of the materials studied and in the interpretation of faunal remains and lithics as testifying to a hominin presence at the site. In actual fact, any reference to the Untermassfeld site as an archaeological one is unwarranted. Furthermore, it is not the only European Early Pleistocene site where inferred evidence for hominin presence is problematic. The strength of the spatiotemporal patterns of hominin presence and absence depends on the quality of the data points we work with, and database maintenance, including critical evaluation of new sites, is crucial to advance our knowledge of the expansions and contractions of hominin ranges during the Pleistocene.

Latest Early Pleistocene wolf-like canids from the Iberian Peninsula

Several species of the genus Canis (Carnivora: Canidae) have been recorded from the European Early Pleistocene, but the phylogenetic relationships among them and in relation to extant members of this genus are still unclear. This is particularly true for the medium-sized and wolf-like extinct species Canis mosbachensis. It has been considered by many researchers as a descendant of the larger Canis etruscus and as a likely putative ancestor of extant wolves (Canis lupus). Other scholars, in contrast, have advocated instead for a closer relationship between C. mosbachensis and the extinct Canis arnensis, and even a close relationship between C. mosbachensis and C. lupus has been questioned. Here we describe the previously unpublished medium-sized Canis remains from the late Early Pleistocene site of Vallparadís Estació, along with additional new Canis material from the roughly coeval site of Cueva Victoria (both in the Iberian Peninsula), and compare them qualitatively and morphometrically with both extant and extinct species of this genus. The described material most closely resembles in craniodental size and shape the remains from Central and Southern Europe that have been previously assigned to C. mosbachensis, to which they are hence formally attributed. The excellent preservation of the newly described specimens (which include the most complete skull of this taxon) enables the description of features previously unknown for this species, which further support a close phylogenetic link with living wolves. Based on the described material, we review the role played by C. mosbachensis in the evolutionary history of European fossil canids, and conclude that this extinct species is most closely related to C. lupus and other closely-allied species, such as Canis anthus and Canis latrans.

An intriguing middle-sized horse from Coste San Giacomo (Anagni Basin, central Italy)

AbstractVarious species and ecomorphotypes of the genus Equus are recorded in several southern European Early Pleistocene local faunal assemblages (LFAs), though their taxonomy, phylogenetic relationships, and chronological distribution are still a matter of debate. This article aims to increase knowledge on the European pre-Olduvai stenonoid horses by describing and discussing the equid sample from the middle Villafranchian (Gelasian) Italian site of Coste San Giacomo (CSG; Anagni). Although horse remains from CSG are scanty, the morphological traits, dimensions, and proportions of teeth and some limb bones suggest some affinities with middle-sized European stenonoid horses, in particular with Equus senezensis, as supported by statistical analysis. This opens a new window on the possible phylogenetic relationships of the middle-sized, slender middle Villafranchian horses, although some questions about their actual taxonomic rank cannot be firmly answered based on available data. The complex relationship among environment, structure of the accompanying mammalian fauna, and the presence in the Early Pleistocene LFAs of only one equid, large or middle-sized, or even of more horse species with different size is briefly discussed.

Early human occupation of Iberia: the chronological and palaeoclimatic inferences from Vallparadís (Barcelona, Spain)

Vallparadís is one of the best calibrated and most accurately dated archaeological sites from the European Early Pleistocene. Chronological analyses combined with palaeomagnetism, ESR-U/series and OSL, and the biochronology of macro- and micromammals are fully consistent and situate the site just above the upper limit of the Jaramillo subchron. In this article we compare the mandibular first molar (m1) of individual adult specimens of Mimomys savini recovered from level 10 (EVT7) at Vallparadís with specimens from the stratigraphic sequence at Gran Dolina (Atapuerca), Fuente Nueva 3 and Barranco León D (Orce). This comparison allows us to chronostratigraphically relate level 10 at Vallparadís with level TD5 at Gran Dolina and to fix the former's chronology to around 0.98–0.95 Ma (MIS 27) and, therefore, prior to level TD6 in which fossil remains of Homo antecessor were recovered. The chronology of Vallparadís and the set of contemporary palaeoclimatic proxies regarding the Iberian Peninsula strengthen the hypothesis that hominins continuously populated Europe, at least in Iberia, throughout the late Early Pleistocene between the Jaramillo subchron and the Matuyama–Brunhes boundary by overcoming the climatic fluctuations and changes to the landscape that occurred during this period.

Continuity or discontinuity in the European Early Pleistocene human settlement: the Atapuerca evidence

The nature, timing, pattern, favourable circumstances and impediments of the human occupation of the European continent during the Early Pleistocene are hot topics in Quaternary studies. In particular, the problem of the (dis) continuity of the settlement of Europe in this period is an important matter of discussion, which has been approached in the last decade from different points of view. The Gran Dolina (TD) and Sima del Elefante (TE) cave sites in the Sierra de Atapuerca, (Spain) include large and quasi-continuous stratigraphic sequences that stretch back from at least 1.2 million years ago (Ma) to the Matuyama/Brunhes boundary. The archaeological and paleontological record from these sites can help to test different hypotheses about the character of the human settlement in this region and period. Furthermore, the TD6 level has yielded a large collection of human fossil remains attributed to Homo antecessor. According to different geochronological methods, as well as to paleomagnetic and biostratigraphical analyses, these hominins belong to an age range of 0.96–0.80 Ma. Unfortunately, the finding in 2007 of some human fossil remains in the TE9 level, dated to about 1.22 Ma, was not enough to conclude whether H. antecessor had deep roots in the European Early Pleistocene. A set of derived features of H. antecessor shared with both the Neanderthal lineage and modern humans suggests that this species is related, and not far, from the most recent common ancestor (MRCA) of Homo neanderthalensis and Homo sapiens. If we assume that there was a lineal biological relationship between the TE9 and TD6 hominins, we should reconsider many of the conclusions achieved in previous paleontological and genetic studies. In addition, we would be obliged to build a highly complicated paleogeographical scenario for the origin of the MRCA. Although continuity in the settlement of Europe during the entire late Early Pleistocene is not discarded (e.g. in refuge areas), it seems that this Western extreme of Eurasia, and the Iberian Peninsula in particular, was occupied by at least two different hominin populations.

Intergroup cannibalism in the European Early Pleistocene: The range expansion and imbalance of power hypotheses

In this paper, we compare cannibalism in chimpanzees, modern humans, and in archaeological cases with cannibalism inferred from evidence from the Early Pleistocene assemblage of level TD6 of Gran Dolina (Sierra de Atapuerca, Spain). The cannibalism documented in level TD6 mainly involves the consumption of infants and other immature individuals. The human induced modifications on Homo antecessor and deer remains suggest that butchering processes were similar for both taxa, and the remains were discarded on the living floor in the same way. This finding implies that a group of hominins that used the Gran Dolina cave periodically hunted and consumed individuals from another group. However, the age distribution of the cannibalized hominins in the TD6 assemblage is not consistent with that from other cases of exo-cannibalism by human/hominin groups. Instead, it is similar to the age profiles seen in cannibalism associated with intergroup aggression in chimpanzees. For this reason, we use an analogy with chimpanzees to propose that the TD6 hominins mounted low-risk attacks on members of other groups to defend access to resources within their own territories and to try and expand their territories at the expense of neighboring groups.

Earliest evidence for human consumption of tortoises in the European Early Pleistocene from Sima del Elefante, Sierra de Atapuerca, Spain

This research was supported by Ministerio de Educacion y Ciencia Spanish Government Grant CGL2009-12703-C03-01, CGL2009-12703-C03-02, CGL2009-12703-C03-03 and CGL2009-7896, and by Generalitat de Catalunya Grant 2009 SGR 188. Ruth Blasco is beneficiary of FI Grant from Generalitat de Catalunya and financed by European Social Found. The field excavation work was supported by Junta de Castilla y Leon and Fundacion Atapuerca.