Carcass transport decisions in Homo antecessor subsistence strategies

Pleistocene foragers used several prey acquisition and processing strategies. These strategies and their associated decisions are elucidated by taphonomic studies that cover animal transport, modifications by different agents and archaeological remains. Interpretative models of archaeological sites are by necessity based on natural and experimental observations. Ethno-archaeological data shows that several factors influenced decisions about carcass transport from the kill site to the home site. These factors often have little archaeological visibility. Díez et al. (1999) has previously interpreted the general characteristics of the macro-mammal remains from Gran Dolina Level TD6-2 (Sierra de Atapuerca, Burgos, Spain) as the result of anthropic accumulation, in which the anatomical profiles appeared to be the result of selective transport based on the animals’ weight. Recent taphonomic analysis has shown that carcasses with different weights may be subject to similar transport strategies, suggesting that other factors influenced these choices. The hominins that occupied TD6-2 (the TD6-2 hominin group), at least sometimes, transported large carcasses to the cave in their entirety, implying participation by groups of individuals in hunting parties. These individuals delayed their consumption of large amounts of food, instead moving it to Gran Dolina, where it was shared with other group members. These decisions are evidence of social cooperation and food sharing amongst early European hominins.     

Carnivores from the Early Pleistocene hominid-bearing Trinchera Dolina 6 (Sierra de Atapuerca, Spain).

The carnivores from Trinchera Dolina level 6 (TD6) in Sierra de Atapuerca include Ursus sp., Crocuta crocuta, Mustela palerminea, Lynx sp., Canis mosbachensis and Vulpes praeglacialis. Approximately 80 human remains belonging to Homo antecessor were found in the Aurora Stratum (AS): located in the upper part of the TD6 level (TD6, T36-43). The carnivores from AS are taxonomically similar to the rest of the TD6 level, differing only in the occurrence of Lynx sp. The spotted hyaena (Crocuta crocuta) inhabited the Atapuerca Sierra during the Early Pleistocene and up to the Middle Pleistocene, after which it is absent. According to palaeomagnetic, U/Th and ESR results, the fossils from TD6 are dated to the Early Pleistocene, which is the earliest certain occurrence in Europe of Crocuta crocuta. It is associated with Mimomys savini, the arrival of which in Europe can be correlated with the beginning of the Early Biharian. Crocuta crocuta may have arrived during the late Early Pleistocene ("Bavelian complex"), and subsequently invaded the rest of the continent during the Middle and Late Pleistocene. Atapuerca TD6-AS represents the most ancient deposits in Europe where Homo and spotted hyaenas coexisted and, they probably competed ecologically. The carnivores from TD6 are biochronologically consistent with the end of the Early Pleistocene or early Cromerian (corresponding with the Biharian biochron). The timespan could correspond with oxygen isotope stages 19, 20 or 21.     

New foot remains from the Gran Dolina-TD6 Early Pleistocene site (Sierra de Atapuerca, Burgos, Spain)

This paper presents and describes new foot fossils from the species Homo antecessor, found in level TD6 of the site of Gran Dolina (Sierra de Atapuerca, Burgos, Spain). These new fossils consist of an almost complete left talus (ATD6-95) and the proximal three-quarters of a right fourth metatarsal (ATD6-124). The talus ATD6-95 is tentatively assigned to Hominin 10 of the TD6 sample, an adult male specimen with which the second metatarsal ATD6-70+107 (already published) is also tentatively associated. Analysis of these fossils and other postcranial remains has made possible to estimate a stature similar to those of the specimens from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain). The morphology of the TD6 metatarsals does not differ significantly from that of modern humans, Neanderthals and the specimens from Sima de los Huesos. Talus ATD6-95, however, differs from the rest of the comparative samples in being long and high, having a long and wide trochlea, and displaying a proportionally short neck.     

Human cannibalism in the Early Pleistocene of Europe (Gran Dolina, Sierra de Atapuerca, Burgos, Spain).

Human remains belonging to at least six individuals were found in an exploratory excavation made at the site of Gran Dolina (Sierra de Atapuerca, Burgos, Spain). These remains were recovered from the Aurora Stratum of Unit TD6. This stratum has a thickness of approximately 30 cm. The area of the exploratory excavation is about 7 m(2). According to palaeomagnetic analyses, Unit TD6 shows reversed polarity, which is considered to belong to the Matuyama chron. This unit is immediately below TD7, where the Matuyama-Brunhes boundary has been detected, indicating an age of around 780,000 years BP. There is no specific distribution, treatment, or arrangement of the human remains, which were found randomly mixed with abundant faunal remains and stone tools. Most of the faunal and human fossil bones from the Aurora Stratum have human induced damage. Stone tool cutmarks are frequent, and peeling (a type of fracture similar to bending a fresh twig between the hands) provides a specific breakage pattern together with percussion marks and chopmarks. Both nonhuman and human remains show similar intensive exploitation. Slight differences, however, have been observed between fauna and humans (e.g., peeling frequent in humans, rare in fauna), that appear related to different musculature, weight, and bone structure. The characteristics of this fossil assemblage suggest that it is solely the result of consumptive activities as there is no evidence of ritual or other intention. The possibility of distinguishing between dietary vs. survival cannibalism is discussed here.     

Hand and foot remains from the Gran Dolina Early Pleistocene site (Sierra de Atapuerca, Spain).

We report here the study of the 22 hand and foot remains from the Early Pleistocene level TD6 of the Gran Dolina site at Sierra de Atapuerca (Burgos, Spain) recovered from 1994 to 1996. These remains are paratypes of Homo antecessor. All of the elements are briefly described and compared with other fossil hominids. The capitate has a constricted neck, well developed head, strong attachment for the ligamentum interosseum trapezoid-capitate, a palmarly placed trapezoid facet with a distinctive small dorsal trapezoid facet, a highly curved and oblique orientation of the second metacarpal facet, and a transversally oriented dorsodistal border. A hamate with a moderately projecting and lightly built hamulus; an inferred reduced styloid process on the third metacarpal base; a wide second metacarpal head; and middle phalanges with well marked insertions for the flexor digitorum superficialis muscle and wide heads. The morphology and dimensions of the pedal remains from TD6 are very similar to modern humans; but the base, proximal articular surface and shafts of the proximal hallucal phalanges are more rounded and the midshaft of the proximal toe phalanx is wider.     

The Atapuerca sites and their contribution to the knowledge of human evolution in Europe

Over the last two decades, the Pleistocene sites of the Sierra de Atapuerca (Spain) have provided two extraordinary assemblages of hominin fossils that have helped refine the evolutionary story of the genus Homo in Europe. The TD6 level of the Gran Dolina site has yielded about one hundred remains belonging to a minimum of six individuals of the species Homo antecessor. These fossils, dated to the end of the Lower Pleistocene (800 kyr), provide the earliest evidence of hominin presence in Western Europe. The origin of these hominins is unknown, but they may represent a speciation event from Homo ergaster/Homo erectus. The TD6 fossils are characterized by a significant increase in cranial capacity as well as the appearance of a “sapiens” pattern of craniofacial architecture. At the Sima de los Huesos site, more than 4,000 human fossils belonging to a minimum of 28 individuals of a Middle Pleistocene population (ca. 500–400 kyr) have been recovered. These hominins document some of the oldest evidence of the European roots of Neanderthals deep in the Middle Pleistocene. Their origin would be the dispersal out of Africa of a hominin group carrying Mode 2 technologies to Europe. Comparative study of the TD6 and Sima de la Huesos hominins suggests a replacement model for the European Lower Pleistocene population of Europe or interbreeding between this population and the new African emigrants.     

The costal skeleton of Homo antecessor: preliminary results

The Lower Pleistocene TD6 level at the Gran Dolina site in the Sierra de Atapuerca (Burgos, Spain) has yielded nine ribs that represent a minimum of three individuals of the species, Homo antecessor. We present a detailed morphological and metric study of these costal elements, including the siding and anatomical position of all of the rib remains. The adult or nearly adult ribs are also metrically compared with other fossil hominins and with modern comparative samples. The costal elements recovered to date from the TD6 level at Gran Dolina can neither confirm nor reject the hypothesis that H. antecessor had a large thorax, similar to that of Neandertals. However, the fragmentary evidence of the H. antecessor thoracic skeleton is not inconsistent with this suggestion based on other skeletal elements, such as clavicles.

Resumen

En el nivel TD6 del Pleistoceno inferior del yacimiento de Gran Dolina, en la Sierra de Atapuerca (Burgos, España) se han recuperado nueve costillas que pertenecen a un mínimo de tres individuos de la especie Homo antecessor. Presentamos un detallado estudio métrico y morfológico incluyendo el lado y la determinación anatómica. Las costillas pertenecientes a individuos adultos o casi adultos también son comparadas métricamente a muestras modernas de comparación y otros homininos fósiles. Basándonos en el registro de costillas de Homo antecessor recuperado hasta el momento no podemos probar ni refutar la hipótesis de que esta especie presentaba un tórax grande similar al de los Neandertales. Sin embargo, el registro de costillas no es inconsistente con la hipótesis de un tórax grande como sugiere la gran longitud de sus clavículas.     

The Pleistocene site of Gran Dolina, Sierra de Atapuerca, Spain: a history of the archaeological investigations.

Gran Dolina is part of an archaeological and paleontological complex located in the Sierra de Atapuerca karstic system (Burgos, Spain). The Trinchera del Ferrocarril sites were discovered as a consequence of the construction of a railway for the transport of minerals at the end of the nineteenth century. The systematic excavation of the upper Gran Dolina levels was initiated in 1981. In 1993, a 6 m(2)biostratigraphic survey pit was started, reaching level TD6 in 1994. This level was excavated during four consecutive years, yielding human fossils, identified as Homo antecessor, in association with lithic and faunal remains, dating to more than 0.78 m.y.a.     

Discontinuity of Human Presence at Atapuerca during the Early Middle Pleistocene: A Matter of Ecological Competition?

Increasing evidence suggests that the European human settlement is older than 1.2 Ma. However, there is a fierce debate about the continuity or discontinuity of the early human settlement of Europe. In particular, evidence of human presence in the interval 0.7−0.5 Ma is scarce in comparison with evidence for the previous and later periods. Here, we present a case study in which the environmental conditions at Sierra de Atapuerca in the early Middle Pleistocene, a period without evidence of human presence, are compared with the conditions in the previous period, for which a relatively intense human occupation is documented. With this objective in mind, the available resources for a human population and the intensity of competition between secondary consumers during the two periods are compared using a mathematical model. The Gran Dolina site TD8 level, dated to 0.7−0.6 Ma, is taken as representative of the period during which Atapuerca was apparently not occupied by humans. Conditions at TD8 are compared with those of the previous period, represented by the TD6-2 level, which has yielded abundant evidence of intense human occupation. The results show that survival opportunities for a hypothetical human population were lower at TD8 than they were at TD6-2. Increased resource competition between secondary consumers arises as a possible explanation for the absence of human occupation at Atapuerca in the early Middle Pleistocene.     

Long-term climate record inferred from early-middle Pleistocene amphibian and squamate reptile assemblages at the Gran Dolina Cave, Atapuerca, Spain

The Gran Dolina cave site is famous for having delivered some of the oldest hominin remains of Western Europe (Homo antecessor, ca. 960 ka). Moreover, the evidence of lithic industries throughout the long vertical section suggests occupation on the part of hominins from the latest early Pleistocene (levels TD3/4, TD5, and TD6) to the late middle Pleistocene (level TD10). The Gran Dolina Sondeo Sur (TDS) has furnished a great number of small-vertebrate remains; among them some 40,000 bones are attributed to amphibians and squamates. Although they do not differ specifically from the extant herpetofauna of the Iberian Peninsula, the overlap of their current distribution areas (= mutual climatic range method) in Spain can provide mean annual temperatures (MAT), the mean temperatures of the coldest (MTC) and warmest (MTW) months, and mean annual precipitation (MAP) estimations for each sub-level, and their change can be studied throughout the sequence. Results from the squamate and amphibian study indicate that during hominin occupation the MAT (10-13 °C) was always slightly warmer than at present in the vicinity of the Gran Dolina Cave, and the MAP (800-1000 mm) was greater than today in the Burgos area. Climatic differences between “glacial” and “interglacial” phases are poorly marked. Summer temperatures (MTW) show stronger oscillations than winter temperatures (MTC), but seasonality remains almost unchanged throughout the sequence. These results are compared with those for large mammals, small mammals, and pollen analysis, giving a scenario for the palaeoclimatic conditions that occurred during the early to middle Pleistocene in Atapuerca, and hence a scenario for the hominins that once lived in the Sierra de Atapuerca.     

Updated Atapuerca biostratigraphy: Small-mammal distribution and its implications for the biochronology of the Quaternary in Spain

The Sierra de Atapuerca (Burgos, Spain) contains the most comprehensive archaeological and paleontological evidence of human evolution in Europe during the Quaternary. The time scale based on the microfossil content of the sedimentary infillings of the caves of Atapuerca has been used to attribute relative ages to the various paleontological and archaeological levels and correlate amongst the different sites. Microfossils are particularly significant as they are the dominant constituents of the caves from the Pleistocene to the Holocene of the Sierra de Atapuerca. The microfossils of the Atapuerca sites are mainly composed of isolated bones and teeth from small vertebrates, and the hard tissue of the vertebrate skeleton can be taxonomically classified. In revisiting the distribution and the classification of the small-mammal taxa along five of the main stratigraphic sequences from Atapuerca (Sima del Elefante, Gran Dolina, Galería-Zarpazos, Sima de los Huesos, and Portalón), we apply a new method, the “Unitary Associations” method, in order to refine the existing biozones, adding five new Faunal Units, and interpret their relative age. Finally, we correlate Atapuerca with other Pleistocene sites of Europe.     

Paleoecological and microenvironmental aspects of the first European hominids inferred from the taphonomy of small mammals (Sima del Elefante,Sierra de Atapuerca,Spain)

Unit TE9 of the Sima del Elefante (Sierra de Atapuerca, Burgos, Spain), where the remains of Homo sp. have been discovered (1.2–1.3 Ma), is also a level rich in small mammals. The taphonomic study of these small vertebrates sheds light on the landscape that provided the setting for the activities of these early hominids and allows us to describe what the cave was like during the formation of the level. Small mammal predators identified in the study indicate that during this period the Sierra de Atapuerca was part of a large biome consisting of semi-open riparian forests with meadows nearby. Postdepositional alterations reveal that the production of fossils took place outside the cave, being transported inside by water currents. During the formation of TE9, the cave presented conditions of high humidity, which made it difficult for hominids to establish occupations inside, although these conditions became somewhat less severe in TE9c, the sublevel where human remains are found.     

A very diverse amphibian and reptile assemblage from the late Middle Pleistocene of the Sierra de Atapuerca (Sima del Elefante,Burgos, Northwestern Spain)

Among the Pleistocene localities of the Sierra de Atapuerca, the site called Sima del Elefante is famous for having delivered in its lower level (TE-LRU) some hominid remains currently considered the oldest in Western Europe (about 1.2 Ma). In its upper part (TE-URU), this site has fossiliferous levels pertaining to the late Middle Pleistocene (350-250 ka). Here we describe for the first time the amphibians and reptiles of these upper levels (TE18 and TE19), which have proved to be one of the richest assemblages of all the localities of the Sierra de Atapuerca. The faunal list is composed of 18 taxa made up of urodeles (Salamandra salamandra and Lissotriton helveticus), anurans (Discoglossus sp., Alytes sp. Pelobates cultripes, Pelodytes punctatus, Bufo bufo, Bufo calamita, Hyla arborea and cf. Rana sp.), a terrestrial tortoise (Testudo s.l.), lizards (Lacerta s.l., Podarcis sp. and Anguis fragilis) and snakes (Natrix natrix, Natrix maura, Coronella cf. girondica and Vipera latasti). For the first time, the genera Lissotriton, Discoglossus and Podarcis are mentioned in the Pleistocene localities of the Sierra de Atapuerca. TE19 also corresponds to the earliest mention for L. helveticus in the Iberian Peninsula. With the exception of tortoises, all these species are currently present in the province of Burgos, and thus suggests that the herpetofauna has a modern aspect since the Middle Pleistocene. This association of amphibians and reptiles suggests a slightly warmer climate than the current one, but may correspond to a “cold and dry” period if compared with other Pleistocene localities of the Sierra de Atapuerca. The landscape was probably composed of a gallery forest along a quiet water river within a Mediterranean environment alternating laterally between dry meadows, rocky or stony areas and open scrubland.     

Étude analytique d’une clavicule complète de subadulte d’Homo antecessor (site de Gran Dolina, Sierra d’Atapuerca, Burgos, Espagne)

This study reports on the skeletal remains of an infant clavicle - specimen ATD6-37 - belonging to the Homo antecessor species, unearthed at Lower Pleistocene level TD6 of the Gran Dolina site (Sierra de Atapuerca). Studied alongside a further adult specimen - ATD6-50 -, they provide us with significant information on two key paleobiological aspects of these early humans: body shape and development. Based on the analytical results, the paper proposes a more accurate proportional method for determining age at death is applied to the fossilized infant clavicle under study. It goes on to hypothesize on postcranial growth and body shape and discusses morphological patterns and age at death of these early humans through comparisons with a wide range of infant dental samples and clavicular specimens in early and modern humans.     

人属先驱种的系统位置——头面骨关键性状的比较研究

人属先驱种(Homo antecessor),是根据西班牙阿塔普尔卡Gran Dolina洞穴堆积中发现的人类化石厘定的。出产人类化石的地层年代测定距今约90万年到78万年之间。之所以定为先驱种,是因为最初研究者认为这个西班牙的古人类群是较晚的海德堡人(H. heidelbergensis)和尼安德特人的共同祖先(H. neanderthalensis),而在非洲和欧洲发现的海德堡人则已被认为是现代人的直接祖先。目前对先驱种的争论还没有明确的结论。之所以会形成这种局面,与"先驱种"模式化石的生物学年龄和保存状况有极大的关系。本文对当初厘定"先驱种"的头骨和面骨的11个特定性状做了一个系统的回顾,尤其是和各类南方古猿(Australopithecus和Praeanthropus),最早的人属成员能人种(H. habilis)和鲁道夫种(H. rudolfensis),人属匠人种(H. ergaster),以及人属直立人种(H. erectus)面部梨状孔周边和上颌骨功能区的特征的比较研究。这11个头面骨的特征包括:1)眶下骨骼的冠状面朝向、2)犬齿窝的存在、3)上颌骨下缘外侧端向下的转折切迹、4)弧状颧骨下缘、5)梨状孔外侧上颌骨的转折和朝向、6)梨状孔外缘和颧骨起点的相对位置、7)梨状孔下缘鼻棘和侧棘的位置和组合形态、8)颧骨和上颌骨结节、9)类现代人的骨表面重塑模式、10)突出的鼻骨、和11)颞骨的上凸的颞鳞上缘。结果表明"先驱种"的头面骨的定性特征没有特异性,其组合与周口店的直立人面部基本形态也非常相似,因而作者倾向于认为"先驱种"就是直立人种的欧洲变异群体。所谓"人属先驱种"可能只是广义的人属直立人种(H. erectus sensu lato)欧洲代表。目前的化石材料显示欧洲和亚洲的直立人种是非洲匠人种的演变而来。     

L’approvisionnement en matières premières lithiques au Pléistocène inférieur et moyen dans la Sierra de Atapuerca, Burgos (Espagne)

This paper summarizes the latest studies on lithic raw material raising that are currently undertaken within the Sierra de Atapuerca research project. During the past two years, several works have provided new petrographical and technological data, contributing to the knowledge of the kinds of materials exploited by the Atapuerca hominids, as well as the areas where those materials were raised. The sites yielding this information are Gran Dolina (Lower and Middle Pleistocene) and Galeria (Middle Pleistocene), both located in the Trinchera del Ferrocarril archeopaleontological complex.     

Axial and appendicular skeleton of Homo antecessor.

The human trunk and limb bones recovered from the Gran Dolina site, in the Sierra de Atapuerca (Burgos, Spain) are studied. All these fossils were excavated at the level called TD6 between 1994 and 1995 and have been dated in excess of 780,000 years ago. These remains have been recently attributed to a new Homo species named Homo antecessor. Axial (vertebrae and ribs) and part of the appendicular (clavicles, radii, femur and patellae) skeleton are studied here. Hand and foot bones have been studied elsewhere (Lorenzo et al., 1999). Four is the minimum number of individuals represented by the postcranial remains recovered up to now. All elements are briefly described anatomically, measured and compared with other fossil hominids and modern humans in order to establish, as far as possible, what postcranial morphology characterized this new species of our genus.The H. antecessor postcrania, generally, display a set of morphological traits that are more similar to modern humans than to the Middle and Upper Pleistocene European hominids. Our results do not contradict the previous phylogenetic analysis, i.e., that H. antecessor represents the last common ancestor for H. sapiens (modern humans) and H. neanderthalensis (Neandertals).     

Climatic and environmental conditions from the Neolithic to the Bronze Age (7000–3000 BP) in the Iberian Peninsula assessed using small-mammal assemblages

We have analyzed the palaeoenvironment and palaeoclimate of the beginning of the Holocene in the Iberian Peninsula on the basis of the small-mammal assemblages from three sites within the geography of the Iberian Peninsula: Mirador (Sierra de Atapuerca, Burgos), Colomera (Sant Esteve de la Sarga, Lleida) and Valdavara-1 (Becerreá, Lugo). These associations reveal that the palaeoenvironment was more humid than today in the sites under study, and the landscape was dominated by woodland and woodland margins in all the studied layers. Further, the climatic conditions were stable, but with winters colder than at present, above all in the Mediterranean area. Finally, our data have been compared with other environmental and climatic proxies, showing that human activities exerted little impact on the palaeoenvironmental conditions that occurred from 7000 to 3000 BP in the Iberian Peninsula.