Gloria Cuenca-Bescós

The first hominin of Europe

The earliest hominin occupation of Europe is one of the most debated topics in palaeoanthropology. However, the purportedly oldest of the Early Pleistocene sites in Eurasia lack precise age control and contain stone tools rather than human fossil remains. Here we report the discovery of a human mandible associated with an assemblage of Mode 1 lithic tools and faunal remains bearing traces of hominin processing, in stratigraphic level TE9 at the site of the Sima del Elefante, Atapuerca, Spain. Level TE9 has been dated to the Early Pleistocene (approximately 1.2–1.1 Myr), based on a combination of palaeomagnetism, cosmogenic nuclides and biostratigraphy. The Sima del Elefante site thus emerges as the oldest, most accurately dated record of human occupation in Europe, to our knowledge. The study of the human mandible suggests that the first settlement of Western Europe could be related to an early demographic expansion out of Africa. The new evidence, with previous findings in other Atapuerca sites (level TD6 from Gran Dolina), also suggests that a speciation event occurred in this extreme area of the Eurasian continent during the Early Pleistocene, initiating the hominin lineage represented by the TE9 and TD6 hominins.

Lower Pleistocene hominids and artifacts from Atapuerca-TD6 (Spain)

Human remains dating to more than 780,000 years ago are associated with a rich faunal and lithic assemblage in the Pleistocene cave site of Gran Dolina (TD), Sierra de Atapuerca, Burgos, Spain. The micromammal species represent the late Biharian (Mimomys savini zone), and the lithic objects represent pre-Acheulean technology (Mode 1) and comes from the TD6 level below the Matuyama-Brunhes boundary. The Gran Dolina hominid fossils cannot be comfortably accommodated in any of the defined Homo species. They could be considered a primitive form of Homo heidelbergensis, but a new species might be named in the future if the sample is enlarged. The new human fossil evidence demonstrates that Western Europe was settled at least since the late early Pleistocene.

Neandertal roots: Cranial and chronological evidence from Sima de los Huesos

Neandertal ancestors from Pleistocene Spain The Sima de los Huesos site in Atapuerca, northern Spain, is a rich source of fossil hominin specimens. The site has now yielded further skull specimens that illuminate patterns of human evolution in Europe nearly half a million years ago. Arsuaga et al. studied 17 crania, including 7 that are new specimens and 6 that are more complete than before (see the Perspective by Hublin). This assemblage of specimens reveals the cranial, facial, and dental features of the Atapuerca hominins, which allows more precise evolutionary positioning of these Neandertal ancestors. Science, this issue p. 1358; see also p. 1338 Seventeen skulls from at least 430 thousand years ago illuminate hominin evolutionary patterns in Pleistocene Europe. [Also see Perspective by Hublin] Seventeen Middle Pleistocene crania from the Sima de los Huesos site (Atapuerca, Spain) are analyzed, including seven new specimens. This sample makes it possible to thoroughly characterize a Middle Pleistocene hominin paleodeme and to address hypotheses about the origin and evolution of the Neandertals. Using a variety of techniques, the hominin-bearing layer could be reassigned to a period around 430,000 years ago. The sample shows a consistent morphological pattern with derived Neandertal features present in the face and anterior vault, many of which are related to the masticatory apparatus. This suggests that facial modification was the first step in the evolution of the Neandertal lineage, pointing to a mosaic pattern of evolution, with different anatomical and functional modules evolving at different rates.

A New Sauropod: Tastavinsaurus Sanzi Gen. Et Sp. Nov. from the Early Cretaceous (Aptian) of Spain

Abstract The new sauropod dinosaur Tastavinsaurus sanzi, gen. et sp. nov., from the early Aptian of Spain is described. The holotype is a partially articulated skeleton of an adult individual recovered from the Arsis-1 site in Peñarroya de Tastavins (Teruel) at the base of the marine Xert Formation. It is one of the most complete and best-preserved sauropod dinosaur skeletons from the European Early Cretaceous. The fossil remains comprise the three caudalmost thoracic vertebrae, part of a fourth, nine thoracic rib fragments, sacrum, 25 caudal vertebrae, 21 chevrons, both ilia, pubes, ischia and femora, right tibia, right fibula, six metatarsals, and seven pedal phalanges (including four unguals). The new taxon is defined by 19 autapomorphies. In our cladistic analysis, Tastavinsaurus is the sister-taxon of the North American Venenosaurus within Titanosauriformes, which includes the Brachiosauridae, Somphospondyli, and Titanosauria. The new taxon provides new information about the diversity of non-brachiosaurid titanosauriforms during the Early Cretaceous in Europe and paleobiogeographic relationships between Europe and North America.

Postcranial morphology of the middle Pleistocene humans from Sima de los Huesos, Spain

Significance The middle Pleistocene Sima de los Huesos (SH) fossil collection provides the rare opportunity to thoroughly characterize the postcranial skeleton in a fossil population, comparable only to that obtained in the study of the Neandertal hypodigm and recent (and fossil) modern humans. The SH paleodeme can be characterized as relatively tall, wide, and muscular individuals, who are less encephalized than both Neandertals and modern humans. Some (but not all) Neandertal derived traits are present, which phylogenetically links this population with Neandertals. Thus, the full suite of Neandertal features did not arise all at once, and the evolution of the postcranial skeleton could be characterized as following a mosaic pattern. Current knowledge of the evolution of the postcranial skeleton in the genus Homo is hampered by a geographically and chronologically scattered fossil record. Here we present a complete characterization of the postcranium of the middle Pleistocene paleodeme from the Sima de los Huesos (SH) and its paleobiological implications. The SH hominins show the following: (i) wide bodies, a plesiomorphic character in the genus Homo inherited from their early hominin ancestors; (ii) statures that can be found in modern human middle-latitude populations that first appeared 1.6–1.5 Mya; and (iii) large femoral heads in some individuals, a trait that first appeared during the middle Pleistocene in Africa and Europe. The intrapopulational size variation in SH shows that the level of dimorphism was similar to modern humans (MH), but the SH hominins were less encephalized than Neandertals. SH shares many postcranial anatomical features with Neandertals. Although most of these features appear to be either plesiomorphic retentions or are of uncertain phylogenetic polarity, a few represent Neandertal apomorphies. Nevertheless, the full suite of Neandertal-derived features is not yet present in the SH population. The postcranial evidence is consistent with the hypothesis based on the cranial morphology that the SH hominins are a sister group to the later Neandertals. Comparison of the SH postcranial skeleton to other hominins suggests that the evolution of the postcranium occurred in a mosaic mode, both at a general and at a detailed level.

Chronological, environmental, and climatic precisions on the Neanderthal site of the Cova del Gegant (Sitges, Barcelona, Spain).

a Ins titu t de Paleoecologia Humana i Evolucio SociaL Area de Prehistiiria de la Universitat Rovira i Virgili. Pla(a Imperial Tarraeo, 1. E-43005 Tarragona, Spain b Departamento de Ciencias de la T ierra, Area de Paleontolog[a de la Universidad de Zaragoza. Cl Pedro Cerbuna, 12. E-50009 Zaragoza, Spain C Centro de Investigaci6n (UCM-ISCIIJ) sobre la Evoluci6n, y Comportamiento Humanos, e/Sinesio Delgado, 4 (Pabe1l6n 14), E-28029 Madrid, Spain cl Dpto. Paleontologfa, Faeultad de Ciencias Geol6gicas, Universidad Complutense de Madrid, Ciudad Universitaria, E-28040 Madrid, Spain

Small-mammals from the Middle Pleistocene layers of the Sima del Elefante (Sierra de Atapuerca, Burgos, northwestern Spain)

The Sima del Elefante site, located in the Sierra de Atapuerca (Burgos, Spain), is an important Pleistocene archaeopalaeontological locality that has been excavated every year since 1996. At least two main infill phases have been identified: a first (TELRU) early Pleistocene phase that has provided a rich faunal assemblage, various stone tools and the earliest human remains from western Europe; and a second phase (TEURU) attributed to the Middle Pleistocene. In this paper, for the first time we present a description of the TEURU small-mammal assemblage and its subsequent biochronological, palaeoenvironmental and palaeoclimatic implications. The small-mammal assemblage is highly diverse and comprises at least 17 species: 3 insectivores (cf. Erinaceus sp., Crocidura sp. and Sorex sp.); 4 chiropters (Rhinolophus ferrumequinum, Rhinolophus gr. euryale-mehelyi, Myotis gr. myotisoxygnathus and Miniopteurs schreibersii); 9 rodents (Myodes glareolus, Microtus arvalis, Microtus agrestis, Iberomys brecciensis, Terricola cf. atapuerquensis, Arvicola sp., Apodemus sylvaticus, Eliomys quercinus and Allocricetus bursae) and 1 lagomorph (Oryctolagus sp.). Such an association suggests a late Middle Pleistocene age (ca. 250-350 ka), a patchy landscape dominated by humid meadows and woodland areas, and mild climatic conditions. These results are compared with other proxies, such as the herpetofauna, malacofauna, large-mammals and charcoals, providing a new scenario for the climatic and environmental conditions that prevailed during the latest Middle Pleistocene in the Sierra de Atapuerca. 2

Environmental change across the Early-Middle Pleistocene transition: small mammalian evidence from the Trinchera Dolina cave, Atapuerca, Spain

Abstract The sites of Atapuerca are well known for their hominin-bearing localities of Early to Middle Pleistocene age. Trinchera Dolina yields western Europe’s oldest fossil hominin, the 850 ka Homo antecessor; and at Sima de los Huesos there occurs the best collection of the Middle Pleistocene hominin Homo heidelbergensis. However, small mammals (Rodentia, Insectivora, Chiroptera and Lagomorpha) represent the largest number of species and greatest fossil abundance in the deposits of the Atapuerca cave complex. The small mammals are grouped by habitat categories, but because the ecology of mammals is complex, we base our study on the evolution of these categories throughout the sequence. The assemblage distribution at the Trinchera Dolina site reflects landscape and environmental changes during the past million years. Shifts between woodland, open land and moorland are indicated by the relative abundance of species based on diagnostic elements such as first lower molars. These data are combined with the distribution of large mammals, pollen, sediments and geological context of the Trinchera Dolina site. From this interdisciplinary approach, we conclude that the first hominins from western Europe (Homo antecessor) lived during a warm, wet and wooded interval, probably corresponding to Marine Isotope Stages 21 to 19. The transition from Early to Middle Pleistocene at Atapuerca is characterized by palaeoenvironmental changes recorded between levels TD5–6 and TD8–10, respectively. The general opening of the landscape at the beginning of the Middle Pleistocene could have favoured the dispersal of hominins (Homo heidelbergensis) across western Europe.

FIRST EVIDENCE OF THE SORICIDAE (MAMMALIA) ASORICULUS GIBBERODON (PETÉNYI, 1864) IN THE PLEISTOCENE OF NORTH IBERIA

The paper describes thirteen specimens of the extinct red-toothed shrew Asoriculus gibberodon (Petenyi, 1864) from the lower levels (8 to 14) of the Sima del Elefante cave (TE Lower Red Unit). These are the first bona fide examples of this taxon in the north of the Iberian peninsula. The Lower Red Unit of the Sima del Elefante dates from the early Pleistocene and contains evidence of the very first human activity in the European continent, in the form of lithic tools and large mammal bones with cutmarks. After a detailed discussion of systematic considerations, we focus on the palaeoecological and palaeobiogeographical implications of the presence of this species in the Sima del Elefantei¦s Lower Red Unit. In palaeoecological terms, as a member of the Neomyini, A. gibberodon would have had a mainly aquatic life and is taken as an indicator of humid environments with a permanent green covering. As for its palaeobiogeographical implications, on the basis of its geographical distribution and morphometric particularities we have reconstructed the biogeography of the genus Asoriculus from its very first record at the end of the Miocene to its extinction shortly before the beginning of the middle Pleistocene.

First evidence of poisonous shrews with an envenomation apparatus

Herein, we report evidence of an envenomation apparatus (EA) in two different species of extinct “giant” shrews, Beremendia and an indeterminate soricine (Mammalia, Eulipotyphla, Soricidae), documented by very well preserved fossil specimens recovered from two Early Pleistocene cave deposits of the Sierra de Atapuerca in Burgos, Spain. The two soricine taxa from Atapuerca have evolved specialized teeth as EAs, which differ from those of recently reported mammals of the Paleocene age, being more similar to the ones described in the modern Solenodon. This discovery reveals the first instance of shrews possessing what appears to be an EA, an evolutionary adaptation that, in these species, was probably related to an increase in body mass and hunting of a larger-sized prey. The Atapuerca specimens would have a highly specialized EA, one of the very few reported for an extinct or living mammal of any time. In addition to the presence of a gutter-like groove along the medial side of the crown of the lower incisors, these two species also present stout jaws and a modified mandibular symphysis with a conspicuous cavity, which in life would likely contain large amounts of connective tissue. The strong mandible architecture of these large shrews would be, in this way, reinforced by a more immovable symphysis, increasing the bite force exerted over a potential prey. This adaptation, together with the grooved incisors, would ensure a rapid and efficient transmission of the poisonous saliva to paralyze relatively large-sized prey.