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.     

How Far into Europe Did Pikas (Lagomorpha: Ochotonidae) Go during the Pleistocene? New Evidence from Central Iberia

This paper reports the first find of pika remains in the Iberian Peninsula, at a site in central Spain. A fragmented mandible of Ochotona cf. pusilla was unearthed from Layer 3 (deposited some 63.4±5.5 ka ago as determined by thermoluminescence) of the Buena Pinta Cave. This record establishes new limits for the genus geographic distribution during the Pleistocene, shifting the previous edge of its known range southwest by some 500 km. It also supports the idea that, even though Europe’s alpine mountain ranges represented a barrier that prevented the dispersal into the south to this and other taxa of small mammals from central and eastern Europe, they were crossed or circumvented at the coldest time intervals of the end of the Middle Pleistocene and of the Late Pleistocene. During those periods both the reduction of the forest cover and the emersion of large areas of the continental shelf due to the drop of the sea level probably provided these species a way to surpass this barrier. The pika mandible was found accompanying the remains of other small mammals adapted to cold climates, indicating the presence of steppe environments in central Iberia during the Late Pleistocene.     

Orientation of DNA Minicircles Balances Density and Topological Complexity in Kinetoplast DNA

Kinetoplast DNA (kDNA), a unique mitochondrial structure common to trypanosomatid parasites, contains thousands of DNA minicircles that are densely packed and can be topologically linked into a chain mail-like network. Experimental data indicate that every minicircle in the network is, on average, singly linked to three other minicircles (i.e., has mean valence 3) before replication and to six minicircles in the late stages of replication. The biophysical factors that determine the topology of the network and its changes during the cell cycle remain unknown. Using a mathematical modeling approach, we previously showed that volume confinement alone can drive the formation of the network and that it induces a linear relationship between mean valence and minicircle density. Our modeling also predicted a minicircle valence two orders of magnitude greater than that observed in kDNA. To determine the factors that contribute to this discrepancy we systematically analyzed the relationship between the topological properties of the network (i.e., minicircle density and mean valence) and its biophysical properties such as DNA bending, electrostatic repulsion, and minicircle relative position and orientation. Significantly, our results showed that most of the discrepancy between the theoretical and experimental observations can be accounted for by the orientation of the minicircles with volume exclusion due to electrostatic interactions and DNA bending playing smaller roles. Our results are in agreement with the three dimensional kDNA organization model, initially proposed by Delain and Riou, in which minicircles are oriented almost perpendicular to the horizontal plane of the kDNA disk. We suggest that while minicircle confinement drives the formation of kDNA networks, it is minicircle orientation that regulates the topological complexity of the network.     

Typing single polymorphic nucleotides in mitochondrial DNA as a way to access Middle Pleistocene DNA

In this study, we have used a technique designed to target short fragments containing informative mitochondrial substitutions to extend the temporal limits of DNA recovery and study the molecular phylogeny of Ursus deningeri. We present a cladistic analysis using DNA recovered from 400 kyr old U. deningeri remains, which demonstrates U. deningeri's relation to Ursus spelaeus. This study extends the limits of recovery from skeletal remains by almost 300 kyr. Plant material from permafrost environments has yielded DNA of this age in earlier studies, and our data suggest that DNA in teeth from cave environments may be equally well preserved.     

Intrapopulational body size variation and cranial capacity variation in middle pleistocene humans: The Sima de los Huesos sample (Sierra de Atapuerca,Spain)

A sexual dimorphism more marked than in living humans has been claimed for European Middle Pleistocene humans, Neandertals and prehistoric modern humans. In this paper, body size and cranial capacity variation are studied in the Sima de los Huesos Middle Pleistocene sample. This is the largest sample of non-modern humans found to date from one single site, and with all skeletal elements represented. Since the techniques available to estimate the degree of sexual dimorphism in small palaeontological samples are all unsatisfactory, we have used the bootstraping method to asses the magnitude of the variation in the Sima de los Huesos sample compared to modern human intrapopulational variation. We analyze size variation without attempting to sex the specimens a priori. Anatomical regions investigated are scapular glenoid fossa; acetabulum; humeral proximal and distal epiphyses; ulnar proximal epiphysis; radial neck; proximal femur; humeral, femoral, ulnar and tibial shaft; lumbosacral joint; patella; calcaneum; and talar trochlea. In the Sima de los Huesos sample only the humeral midshaft perimeter shows an unusual high variation (only when it is expressed by the maximum ratio, not by the coefficient of variation). In spite of that the cranial capacity range at Sima de los Huesos almost spans the rest of the European and African Middle Pleistocene range. The maximum ratio is in the central part of the distribution of modern human samples. Thus, the hypothesis of a greater sexual dimorphism in Middle Pleistocene populations than in modern populations is not supported by either cranial or postcranial evidence from Sima de los Huesos. Am J Phys Anthropol 106:19–33, 1998. © 1998 Wiley-Liss, Inc.     

Cranial discrete traits in the middle pleistocene humans from Sima de los Huesos (Sierra de Atapuerca, Spain). Does hypostosis represent any increase in "ontogenetic stress" along the Neanderthal lineage?

Cranial discrete traits may be regarded as markers of dynamic responses to general and local perturbations of the morphogenetic pattern, particularly when they are viewed and examined in terms of hypostosis vs. hyperostosis. There are indications, in fact, that the variation between these two opposite conditions relates to mechanical stress suffered by the bony structures during early stages of growth and development. In a previous comparison between Neanderthals and modern humans, variable degrees and contrasting distribution patterns of hypostosis were found [Manzi et al. (1996), JHE30: 511-527]. In the present paper, the occurrence, expression and cranial distribution of 20 hypo-hyperostotic traits are examined in the Middle Pleistocene sample from Atapuerca - Sima de los Huesos (Spain), with the principal aim being to test whether or not the degree of cranial hypostosis increases during the evolution of the Neanderthals. Other Middle Pleistocene representatives of the genus Homo (Kabwe and Petralona), the Italian Neanderthals, and a large recent European sample are also considered. A general consistency between the gradual appearance and stabilization of the Neanderthal cranial features and the results of the present analysis is found and is interpreted as an indication that hypostosis does mark the occurrence of "ontogenetic stress". As suggested more than half a century ago by S. Sergi, an increase in "ontogenetic stress" in the Neanderthal lineage could result from the relationship between intracranial pressures and other (heterochronic) effects produced by the growth of a large brain (encephalization) and the ossification of an archaic (platycephalic) cranial vault.     

Les premiers comportements funéraires auraient-ils pris place à Atapuerca, il y a 350 000 ans ?

A lithic tool has appeared for the first time at Sima de los Huesos (Atapuerca, Spain). It is a finely flaked quartzite handaxe, which is associated with the hominid assemblage of a MNI of 27 Homo heidelbergensis, dated more than 350,000 years ago. The particular nature of the deposit involving its taphonomy, paleontology, and technology points to a symbolic meaning both of the tool and the human accumulation.     

Chromosomes are predominantly located randomly with respect to each other in interphase human cells

To test quantitatively whether there are systematic chromosome-chromosome associations within human interphase nuclei, interchanges between all possible heterologous pairs of chromosomes were measured with 24-color whole-chromosome painting (multiplex FISH), after damage to interphase lymphocytes by sparsely ionizing radiation in vitro. An excess of interchanges for a specific chromosome pair would indicate spatial proximity between the chromosomes comprising that pair. The experimental design was such that quite small deviations from randomness (extra pairwise interchanges within a group of chromosomes) would be detectable. The only statistically significant chromosome cluster was a group of five chromosomes previously observed to be preferentially located near the center of the nucleus. However, quantitatively, the overall deviation from randomness within the whole genome was small. Thus, whereas some chromosome-chromosome associations are clearly present, at the whole-chromosomal level, the predominant overall pattern appears to be spatially random.     

Earliest humans in Europe: the age of TD6 Gran Dolina, Atapuerca, Spain

Hominid remains found in 1994 from the stratified Gran Dolina karst-filling at the Atapuerca site in NE Spain were dated to somewhat greater than 780 ka based on palaeomagnetic measurements, making these the oldest known hominids in Europe (sensu stricto). We report new ESR and U-series results on teeth from four levels of the Gran Dolina deposit which confirm the palaeomagnetic evidence, and indicate that TD6 (from which the human remains have been recovered) dates to the end of the Early Pleistocene. The results for the other levels are consistent with estimates based mainly on microfaunal evidence, and suggest that TD8, TD10 and TD11 date to the Middle Pleistocene.     

Cranial and mandibular morphology of Middle Pleistocene cave bears (Ursus deningeri): implications for diet and evolution

ABSTRACT

Deninger’s bears (Ursus deningeri) have been studied less frequently than Ursus spelaeus s.l. Our objective is to present, for the first time, an analysis of the skull shape of U. deningeri.

Bear crania and mandibles were digitised with a Microscribe or CT-scanned and the surface models subsequently landmarked. The landmarks were chosen based on a compromise between functional morphology and sample size.

Results show that U. deningeri and U. spelaeus mandibles display very similar morphologies and allometric trajectories, both to each other and to Ailuropoda melanoleuca. It is inferred that masticatory adaptations to a herbivorous diet were already present in the Middle Pleistocene. U. deningeri displays a cranial morphology that is similar to that of U. spelaeus when comparing all species, but U. deningeri has a relatively narrower and dorsoventrally lower zygomatic arch than U. spelaeus, although the masticatory signal is less strong in the skull.

We observe intraspecific differences between different populations of U. deningeri, which could parallel the genetic diversity found in U. spelaeus. The intraspecific differences found within U. deningeri may be temporal and/or geographical in nature and could be related to the evolution of the Late Pleistocene cave bear, but this hypothesis remains to be tested.