The influence of raw material qualities in the lithic technology of Gran Dolina (Units TD6 and TD10) and Galería (Sierra de Atapuerca,Burgos, Spain): A view from experimental archeology

This paper analyses the qualities of the raw materials used in two Palaeolithic sites (Gran Dolina and Galería) of the Sierra de Atapuerca (Burgos, Spain) during the Lower and Middle Pleistocene, and their influence in the development of knapping. These sites offer a chronological sequence that allows us to study the evolution of lithic technology at a local scale during 1.2 Ma. Combining technological analysis and experimental archaeology has proven to be an excellent tool for the understanding and the interpretation of the qualities of raw materials and their relation with the development of the gestures, methods and techniques.     

Posterior dental size reduction in hominids: The Atapuerca evidence

In order to reassess previous hypotheses concerning dental size reduction of the posterior teeth during Pleistocene human evolution, current fossil dental evidence is examined. This evidence includes the large sample of hominid teeth found in recent excavations (1984–1993) in the Sima de los Huesos Middle Pleistocene cave site of the Sierra de Atapuerca (Burgos, Spain). The lower fourth premolars and molars of the Atapuerca hominids, probably older than 300 Kyr, have dimensions similar to those of modern humans. Further, these hominids share the derived state of other features of the posterior teeth with modern humans, such as a similar relative molar size and frequent absence of the hypoconulid, thus suggesting a possible case of parallelism. We believe that dietary changes allowed size reduction of the posterior teeth during the Middle Pleistocene, and the present evidence suggests that the selective pressures that operated on the size variability of these teeth were less restrictive than what is assumed by previous models of dental reduction. Thus, the causal relationship between tooth size decrease and changes in food-preparation techniques during the Pleistocene should be reconsidered. Moreover, the present evidence indicates that the differential reduction of the molars cannot be explained in terms of restriction of available growth space. The molar crown area measurements of a modern human sample were also investigated. The results of this study, as well as previous similar analyses, suggest that a decrease of the rate of cell proliferation, which affected the later-forming crown regions to a greater extent, may be the biological process responsible for the general and differential dental size reduction that occurred during human evolution. © 1995 Wiley-Liss, Inc.     

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.     

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.     

Pattern of dental development in Hominid XVIII from the Middle Pleistocene Atapuerca-Sima de los Huesos site (Spain)

. We describe the pattern of dental development of Hominid XVIII from the Middle Pleistocene Sima de los Huesos (SH) site of the Sierra de Atapuerca (Burgos, Spain). As expected, this pattern is similar to that of modern humans. A delay of development of the lower and upper canines was observed. In contrast, the relative advanced development of the lower second molars and, especially, the upper and lower third molars is noteworthy. This latter feature seems to be common in Pleistocene hominids, and suggests that the pattern of dental development evolved in the genus Homo during the Pleistocene. In European Middle Pleistocene hominids, this pattern probably was facilitated by the extra space available in the mandible and maxilla for developing teeth.     

Early and Middle Pleistocene hominins from Atapuerca (Spain) show differences in dental developmental patterns

The Bayesian statistical approach considers teeth as forming a developmental module, as opposed to a tooth-by-tooth analysis. This approach has been employed to analyze Upper Pleistocene hominins, including Neandertals and some anatomically modern humans, but never earlier populations. Here, we show its application on five hominins from the TD6.2 level of the Gran Dolina site (Homo antecessor, Early Pleistocene) and the Sima de los Huesos site (Middle Pleistocene) of the Sierra de Atapuerca (Burgos, northern Spain). Our results show an advanced development of the third molars in both populations with respect to modern Homo sapiens. In addition, the Sima de los Huesos hominins differ from H. sapiens and H. antecessor in the relatively advanced development of their second molar. The relative mineralization of I1/M1 in H. antecessor appears to be similar to that of modern humans, as opposed to that of Neandertals, which appear to be unique. These observations, combined with reduced enamel formation times and the advanced development of the third molars, appear to indicate a shorter ontogenetic period in the hominins from Gran Dolina and Sima de los Huesos in comparison to modern human average.     

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 geometric morphometric analysis of hominin upper second and third molars, with particular emphasis on European Pleistocene populations

The study of dental morphology by means of geometric morphometric methods allows for a detailed and quantitative comparison of hominin species that is useful for taxonomic assignment and phylogenetic reconstruction. Upper second and third molars have been studied in a comprehensive sample of Plio- and Pleistocene hominins from African, Asian and European sites in order to complete our analysis of the upper postcanine dentition. Intraspecific variation in these two molars is high, but some interspecific trends can be identified. Both molars exhibit a strong reduction of the distal cusps in recent hominin species, namely European Homo heidelbergensis, Homo neanderthalensis and Homo sapiens, but this reduction shows specific patterns and proportions in the three groups. Second molars tend to show four well developed cusps in earlier hominin species and their morphology is only marginally affected by allometric effects. Third molars can be incipiently reduced in earlier species and they evince a significant allometric component, identified both inter- and intraspecifically. European Middle Pleistocene fossils from Sima de los Huesos (SH) show a very strong reduction of these two molars, even more marked than the reduction observed in Neanderthals and in modern human populations. The highly derived shape of SH molars points to an early acquisition of typical Neanderthal dental traits by pre-Neanderthal populations and to a deviation of this population from mean morphologies of other European Middle Pleistocene groups.     

Sea-surface temperature for left-coiling Neogloboquadrina populations inhabiting the westernmost Mediterranean in the middle Pleistocene and the Pleistocene-Pliocene transition

Using the modern analog methodology applied to planktonic foraminifers, we analyze the relation between the frequency of the coiling type in Neogloboquadrina populations and the sea-surface temperatures (SST) during the middle Pleistocene and the Pleistocene-Pliocene transition in the Alboran Sea (westernmost Mediterranean), close to the Atlantic connection. The results reveal that the present-day positive correlation between the two variables (r = 0.649) is maintained even with a higher coefficient (r = 0.783) in the middle Pleistocene but falls slightly (r = 0.517) in the Pleistocene-Pliocene transition due mainly to a dispersal of the temperatures for the samples bearing predominantly left-coiling Neogloboquadrina. The temperature used as a reference for the coiling change resulted 5-6 °C above its North Atlantic present-day reciprocal, but these thermal differences could be caused by: i) a slight overestimate (1–2 °C) due to the lumping of all neogloboquadrinids into a single variable when SST are estimated; ii) only the warmer range of temperatures for the left-coiling populations would be represented in the samples; and iii) a remarkable warming inside the westernmost Mediterranean during the summer stage mixing cold and warm assemblages in bottom sediments. In addition, these results in combination with those derived from the isotopic analyses (δ¹⁸O) in G. bulloides tests, suggest that during the Pliocene-Pleistocene transition two different populations of left-coiling Neogloboquadrina could have existed with different environmental requirements: one, derived from late Miocene-Pliocene (i.e., left-coiling N. acostaensis group), and another being the ancestor of the modern N. pachyderma (left-coiling). Similarity analyses were achieved in order to locate the position of the core-tops with the assemblages most analogous to those of the fossil samples containing left-coiling Neogloboquadrina. The results reinforce the idea that the presence of these forms in the Mediterranean during the intervals studied would be related mainly to the input of cold waters from the North Atlantic during glacial stages, although it could be secondarily favored by the establishment of upwelling conditions, as in the present-day North Alboran waters.     

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.     

Reconstitution 3D par Computerized-tomography (CT) et endocrâne virtuel du crâne 5 du site de la Sima de Los Huesos (Atapuerca)

The 3d Ct reconstruction and virtual brain endocast of Cranium 5 from the site of “La Sima de los Huesos” (Atapuerca), allows us to get new information to increase the study and knowledge of Homo heidelbergensis population, and to compare this specimen with others in the fossil record, in order to understand the evolutionary process of the brain, focusing on the middle Pleistocene period. Furthermore, we can observe the changes this species (Homo heidelbergensis) has undergone, at least in Sima de los Huesos population making comparative studies with African and Asian middle Pleistocene specimens. We have used the new data to compare European Homo heidelbergensis represented by SH5 with Kabwe, a controversial specimen considered by some authors like the African Homo heidelbergensis representative, in order to establish the similarities and differences between both specimens.     

Timing of clade divergence and discordant estimates of genetic and morphological diversity in the Slender Madtom,Noturus exilis (Ictaluridae)

Phylogeographic relationships, the timing of clade diversification, and the potential for cryptic diversity in the Slender Madtom, Noturus exilis, was investigated using mitochondrial Cyt b, nuclear RAG2, shape analysis, and meristic and pigmentation data. Three well-supported and deeply divergent clades were recovered from analyses of genetic data: Little Red River (White River drainage) clade, Arkansas + Red River (Mississippi River) clade, and a large clade of populations from the rest of the range of the species. Recovered clades showed little to no diagnostic morphological differences, supporting previous hypotheses of morphological conservatism in catfishes, and indicating morphology may commonly underestimate diversity in this group of fishes. The Little Red River clade is the most distinct lineage of N. exilis with 11 POM pores (vs. 10 in other populations) and unique Cyt b haplotypes and RAG 2 alleles. However, treating it as a species separate from N. exilis would imply that the other major clades of N. exilis are more closely related to one another than they are to the Little Red River clade, which was not supported.The UCLN age estimate for Noturus was 23.9 mya (95% HPD: 13.49, 35.43), indicating a late Oligocene to early Miocene origin. The age of N. exilis was estimated as late Miocene at 9.7 mya (95% HPD: 5.32, 14.93). Diversification within the species spanned the late Miocene to mid-Pleistocene. The largest clade of N. exilis, which dates to the late Miocene, includes populations from the unglaciated Eastern and Interior Highlands as well as the previously glaciated Central Lowlands. Diversification of this clade coincides with a drastic drop in sea-level and diversification of other groups of Central Highlands fishes (Centrarchidae and Cyprinidae). Sub-clades dating to the Pleistocene show that northern populations occurring in previously glaciated regions resulted from dispersal from populations in the Ozarks up the Mississippi River following retreat of the Pleistocene glaciers. Pre-Pleistocene vicariance, such as drainage pattern changes of the Mississippi River, also played a prominent role in the history of the species. The incorporation of a temporal estimate of clade diversification revealed that in some instances, phylogeographic breaks shared with other aquatic species were best explained by different or persistent vicariant events through time, rather than a single shared event.     

Stephanorhinus hundsheimensis (Mammalia,Rhinocerotidae) from the late early Pleistocene deposits of the Denizli Basin (Anatolia,Turkey)

Pleistocene rhinoceroses are poorly documented in Turkey where they have been reported only from the late early Pleistocene (1.3–1.1 Ma) travertine deposits of the Denizli Basin. In this work, new rhinoceros remains collected from this basin are assigned to a relatively large-sized Stephanorhinus hundsheimensis on the basis of their morphology and morphometry. The first Turkish record of this species is approximately coeval with the first appearance of S. hundsheimensis in Europe, chronologically referred to the late early Pleistocene, ca. 1.2 Ma. During that time, S. etruscus still survived in Iberian Peninsula, central Italy and Dacian Basin. The presence of two successive evolutionary morphs for S. hundsheimensis during the Pleistocene is not confirmed.     

A human parietal fragment from the late Early Pleistocene Gran Dolina-TD6 cave site,Sierra de Atapuerca,Spain

The TD6-2 level of the Gran Dolina cave site (Sierra de Atapuerca, Burgos, Spain) has yielded an assemblage of about 170 human fossil remains dated to > 800 ka (probably MIS 21) and assigned to the species Homo antecessor. In this study, we describe for the first time a large portion of a parietal bone (ATD6-100/168). The morphology of the fractures on the bones is compatible with a peri-mortem trauma. The superior parietal areas are flat. There is a large parietal foramen and one smaller accessory parietal foramen. Middle meningeal vessels are not particularly developed, but they are distributed in both anterior and posterior districts, with the parietal vasculature originating from the posterior branch. The meningeal vessels show multiple minor connections with the pericranial and diploic vascular systems. The diploe is not particularly developed, and large diploic channels are not detected. The bone is thin when compared with adult fossil humans, and equivalent to juvenile values. All these characters suggest that the parietal ATD6-100/168 probably belonged to a juvenile individual, with plesiomorphic endocranial traits similar to those described for H. ergaster/erectus. The derived temporal, maxillary and dental traits in Homo antecessor and the primitive parietal morphology further point to distinct (mosaic) patterns of morphological evolution of face and braincase.     

Interleukin-6 expression on inflamed rat dental pulp tissue after capped with Trigona sp. propolis from south Sulawesi,Indonesia

Background: Propolis is a natural product of plant resins collected by honeybees from various plant sources. It is used as a remedy in folk medicine since ancient times because of its several biological and pharmacological properties. Recently, propolis has been used by dentist to treat various oral diseases. It was always mentioned as an anti-inflammatory agent. Cytokines are proteins that provide communication between cells and play a critical role in a wide variety of processes. It released from cells in an inflammatory process that active, mediate or potential actions of other cells or tissues. When dental pulp has inflammation, several pro-inflammatory cytokines including Interleukin-6 (IL-6) was released by innate immune cells. Objective: To analyse the expression of IL-6 on inflamed rat dental pulp tissue following application of propolis. Material and methods: Trigona sp. propolis was obtained from Luwu Regency, south Sulawesi Province, Indonesia. Flavonoid and non-flavonoid extracts were purified from propolis using thin layer chromatography. The study was applied on 80 male Sprague Dawley rats, 10–12 weeks of age, divided randomly and equally into 5 groups. Group I, as negative control group was not conducted any treatment. At group II, III, IV and V. A Class I cavity (Black Classification) were made on the occlusal surface of right maxillary first molar. The dental pulp was perforated using dental explorer and allowed in the oral environment for 1 h, after that, Ethanolic Extract Propolis (EEP) (Group II), Extract Flavonoid-Propolis (EFP) (Group III), Extract Non-Flavonoid Propolis (ENFP) (Group IV), or Calcium Hydroxide (Ca(OH)₂) (Group V) were applied on dental pulp. All cavities were then filled with Glass Ionomer Cement as permanent filling. The rats being sacrificed in 6 h, 2 days, 4 days and 7 days. Sample biopsy were obtained, IL-6 expression was detected by using immunohistochemistry method. Data was analyzed statistically using Freidman and Kruskal Wallis tests with significance level of P < 0.05. Results: All agent showed IL-6 expression in inflamed rat dental pulp tissue, and this expression was decreased with the longer of observation time periods. EEP more stronger to decreased IL-6 expression on inflamed rat dental pulp tissue than other agent. There is significant difference (P < 0.05) of IL-6 expression between group I and other groups in 6 h and 2 days but not in 4 and 7 days time periods. Conclusion: Trigona sp. propolis from south Sulawesi, Indonesia could suppressed the expression of IL-6 on inflamed rat dental pulp tissue.     

Tissue proportions and enamel thickness distribution in the early Middle Pleistocene human deciduous molars from Tighenif,Algeria

The present study of three human upper deciduous molars from the early Middle Pleistocene site of Tighenif, Algeria, constitutes the first microtomographic-based endostructural exploration of African fossil teeth likely representative of the Homo heidelbergensis morph. Comparative morphological observations and 2-3D measurements describing subtle tooth organization (crown tissue proportions) and enamel thickness topography (site-specific distribution and global patterning) indicate that their virtual extracted structural signature better fits the modern human, rather than the Neanderthal condition. Accordingly, we predict that the inner structural morphology of the deciduous molars from the Middle Pleistocene western European series better fits the primitive, and not the derived Neanderthal figures.     

Human hyoid bones from the middle Pleistocene site of the Sima de los Huesos (Sierra de Atapuerca, Spain)

This study describes and compares two hyoid bones from the middle Pleistocene site of the Sima de los Huesos in the Sierra de Atapuerca (Spain). The Atapuerca SH hyoids are humanlike in both their morphology and dimensions, and they clearly differ from the hyoid bones of chimpanzees and Australopithecus afarensis. Their comparison with the Neandertal specimens Kebara 2 and SDR-034 makes it possible to begin to approach the question of temporal variation and sexual dimorphism in this bone in fossil humans. The results presented here show that the degree of metric and anatomical variation in the fossil sample was similar in magnitude and kind to living humans. Modern hyoid morphology was present by at least 530 kya and appears to represent a shared derived feature of the modern human and Neandertal evolutionary lineages inherited from their last common ancestor.     

A geometric morphometric analysis of hominin upper premolars. Shape variation and morphological integration

This paper continues the series of articles initiated in 2006 that analyse hominin dental crown morphology by means of geometric morphometric techniques. The detailed study of both upper premolar occlusal morphologies in a comprehensive sample of hominin fossils, including those coming from the Gran Dolina-TD6 and Sima de los Huesos sites from Atapuerca, Spain, complement previous works on lower first and second premolars and upper first molars. A morphological gradient consisting of the change from asymmetric to symmetric upper premolars and a marked reduction of the lingual cusp in recent Homo species has been observed in both premolars. Although percentages of correct classification based on upper premolar morphologies are not very high, significant morphological differences between Neanderthals (and European middle Pleistocene fossils) and modern humans have been identified, especially in upper second premolars. The study of morphological integration between premolar morphologies reveals significant correlations that are weaker between upper premolars than between lower ones and significant correlations between antagonists. These results have important implications for understanding the genetic and functional factors underlying dental phenotypic variation and covariation.