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New pedal remains of Megaladapis and their functional significance.

New remains of Megaladapis from the caves within the Ankarana Range of northern Madagascar and the cave site of Ankilitelo near Toliara in southwestern Madagascar add considerably to the present sample of pedal remains for this genus. Here we describe and analyze the new pedal material and discuss the function of the Megaladapis foot in terms of positional behavior and substrate use. The northern specimens belong to the M. madagascariensis /M. grandidieri group in terms of size and morphology whereas the new southwestern fossils are assigned to M. madagascariensis. The new specimens demonstrate that the small and intermediate sized M. madagascariensis and M. grandidieri were very similar in anatomy and inferred locomotor function, findings that also support the prior suggestion that they belong to a single widespread subgenus (Megaladapis). The new fossils provide the first examples of many pedal elements and present the first opportunity to analyze the whole pedal complex from associated remains. The foot of Megaladapis is distinctive among primates in numerous features. Intrinsic proportions of the hindlimb indicate that the foot is relatively longer than that of any other primate. The first complete calcanei reveal a large and highly modified hindfoot. The calcaneus is reduced distally, indicating an emphasis on climbing over leaping or quadrupedal walking and running. Proximally, a large, medially directed calcaneal tuberosity suggests both a strong inversion component to plantarflexion and a well-developed abductor mechanism and recalls the calcaneal morphology of the larger lorisines in some respects. Talar shape is consistent with considerable tibial rotation during plantarflexion and dorsiflexion. The subtalar joint is designed to emphasize supination/pronation and medial/lateral rotation over proximodistal translation. The distal tarsals are extremely reduced in length, and they form a high transverse arch and a serial tarsus; this configuration promotes inversion/eversion at the transverse tarsal joint. The phalanges are long and moderately curved, and the hallux is very long, robust, and abducted. Pedal morphology suggests that Megaladapis (subgenus Megaladapis) was well adapted to exploit an arboreal environment.(ABSTRACT TRUNCATED)

Animals↗

Particulate versus integrated evolution of the upper body in late pleistocene humans: a test of two models.

Evolutionary biologists are largely polarized in their approaches to integrating microevolutionary and macroevolutionary processes. Neo-Darwinians typically seek to identify population-level selective and genetic processes that culminate in macroevolutionary events. Epigeneticists and structuralists, on the other hand, emphasize developmental constraints on the action of natural selection, and highlight the role of epigenetic shifts in producing evolutionary change in morphology. Accordingly, the ways in which these paradigms view and address morphological contrasts between classes of related organisms differ. These paradigms, although seldomly explicitly stated, emerge in paleoanthropology as well. Considerations of postcranial morphological contrasts between archaic and modern humans typically fall into one of two broad interpretive models. The first derives from the neo-Darwinian perspective and holds that evolution in the postcranial skeleton was largely mosaic (operating in a particulate manner), and that temporal change in specific traits informs us about behavioral shifts or genetic evolution affecting isolated anatomical regions (i.e., adaptive behavioral inferences can be made from comparative studies of individual trait complexes). The alternative model follows from the epigeneticist paradigm and sees change in specific postcranial traits as correlated responses to change in overall body form (involving shifts in regulation of skeletal growth, or selective and developmental responses to broad adaptive shifts). By this view, integration of functional systems both constrains and directs evolution of various traits, and morphological contrasts inform us about overall change in body form related to change in such things as overall growth patterns, climatic adaptation, and technological dependency. These models were tested by confirmatory factor analysis using measures of upper body form and upper limb morphological traits in Eurasian Neandertal and early modern fossils and recent human samples. Results indicate (1) a model of morphological integration fits the data better than a model of no integration, but (2) this integration accounts for less than half of the variance in upper limb traits, suggesting a high degree of tolerance for particulate evolution in the context of an integrated upper body plan. Significant relationships were detected between joint shapes and body size, between humeral shaft shape and body size and chest shape, and between measures of biomechanical efficiency and robusticity. The observed morphological differences between late archaic and early modern humans reflect particulate evolution in the context of constraints imposed by genetic and morphological integration. While particulate approaches to interpreting the fossil record appear to be justified, attention must also be paid to delineating the nature and extent of morphological integration and its role in both constraining and producing observed patterns of variation between groups. Confirmatory factor analysis provides a means of examining trait covariance matrices, and serves as a useful method of identifying patterns of integration in morphology.

Biological Evolution↗

Developmental age and taxonomic affinity of the Mojokerto child, Java, Indonesia.

An increasing number of claims place hominids outside Africa and deep in Southeast Asia at about the same time that Homo erectus first appears in Africa. The most complete of the early specimens is the partial child's calvaria from Mojokerto (Perning I), Java, Indonesia. Discovered in 1936, the child has been assigned to Australopithecus and multiple species of Homo, including H. modjokertensis, and given developmental ages ranging from 1-8 years. This study systematically assesses Mojokerto relative to modern human and fossil hominid growth series and relative to adult fossil hominids. Cranial base and vault comparisons between Mojokerto and H. sapiens sapiens (Hss) (n = 56), Neandertal (n = 4), and H. erectus (n = 4) juveniles suggest a developmental age range between 4 and 6 years. This range is based in part on new standards for assessing the relative development of the glenoid fossa. Regression analyses of vault arcs and chords indicate that H. erectus juveniles have more rounded frontals and less angulated occipitals than their adult counterparts, whereas Hss juveniles do not show these differences relative to adults. The growth of the cranial superstructures and face appear critical to creating differences in vault contours between H. erectus and Hss. In comparison with adult H. erectus and early Homo (n = 27) and adult Hss (n = 179), the Mojokerto child is best considered a juvenile H. erectus on the basis of synapomorphies of the cranial vault, particularly a metopic eminence and occipital torus, as well as a suite of characters that describe but do not define H. erectus, including obelion depression, supratoral gutter, postorbital constriction, mastoid fissure, lack of sphenoid contribution to glenoid fossa, and length and breadth ratios of the temporomandibular joint. Mojokerto is similar to other juvenile H. erectus in the degree of development of its cranial superstructures and its vault contours relative to adult Indonesian specimens. The synapomorphies which Mojokerto shares with H. erectus are often considered autapomorphies of Asian H. erectus and confirm the early establishment and long-term continuity of the Asian H. erectus bauplan. This continuity does not, however, necessarily reflect on the pattern of origin of modern humans in the region.

Adolescent↗

Brief communication: evidence of pathology on the frontal bone from Gongwangling.

The hominid fossil from Gongwangling (Lantian) is well known and described (Woo, [1965] Scientia Sinca 14:1032-1036; Woo [1966] Curr. Anthropol. 7:83-86; Wu and Dong [1985] in R Wu and JW Olsen (eds.): Palaeoanthropology and Paleolithic Archaeology in the People's Republic of China [New York: Academic Press, pp. 79-89]; Wu and Poirier [1995] Human Evolution in China: A Metric Description of the Fossils and a Review of the Sites [Oxford: Oxford University Press]). However, evidence of pathology on the frontal bone has been previously unreported. Two lesions occur on the right supraorbital region that can be distinguished from marks of erosion prevalent on this specimen. These are discrete and irregularly shaped, with evidence of secondary bone formation surrounding them. The cause of the condition is unclear. Possibilities include trauma or abscess from an unspecified infection.

Abscess↗

Phalangeal morphology of the paromomyidae (?primates, plesiadapiformes): the evidence for gliding behavior reconsidered.

A comparative morphometric analysis of isolated proximal and intermediate phalanges attributed to the paromomyids Ignacius graybullianus and Phenacolemur simonsi was undertaken to test the hypothesis that these fossil phalanges exhibit evidence of a dermopteran-like interdigital patagium. Linear dimensions were collected for the fossil phalanges and a comparative sample of associated proximal and intermediate phalanges representing extant tree squirrels, tree shrews, dermopterans (colugos), gliding rodents and marsupials, and prosimian primates. Quantitative data indicate that the proximal and intermediate phalanges of paromomyids are most similar in their overall shape to those of the dermopteran Cynocephalus. The proximal phalanges of paromomyids and colugos possess well-developed flexor sheath ridges and broad, high shafts, whereas the intermediate phalanges of these taxa are most similar to one another in their trochlear morphology. Discriminant analysis indicates that all of the paromomyid intermediate phalanges resemble those from colugo toes more so than those from colugo fingers. Moreover, the relative length and midshaft proportions of both the proximal and intermediate phalanges of paromomyids closely resemble those of several squirrels that lack an interdigital patagium. The following conclusions are drawn from this study: 1) paromomyids share a number of derived phalangeal features with modern dermopterans that may be indicative of a phylogenetic relationship between them, 2) existing intermediate phalanges of paromomyids are inconsistent with the "mitten gliding" hypothesis because they do not possess the distinctive length and midshaft proportions characteristic of colugo manual intermediate phalanges, and 3) paromomyids share with colugos and the scaly-tailed squirrel Anomalurus several aspects of phalangeal morphology functionally related to frequent vertical clinging and climbing on large-diameter arboreal supports.

Animals↗

A modern human humerus from the early aurignacian of Vogelherdhöhle (Stetten, Germany).

Implicit in much of the discussion of the cultural and population biological dynamics of modern human origins in Europe is the assumption that the Aurignacian, from its very start, was made by fully modern humans. The veracity of this assumption has been challenged in recent years by the association of Neandertal skeletal remains with a possibly Aurignacian assemblage at Vindija Cave (Croatia) and the association of Neandertals with distinctly Upper Paleolithic (but non-Aurignacian) assemblages at Arcy-sur-Cure and St. C¿esaire (France). Ideally we need human fossil material that can be confidently assigned to the early Aurignacian to resolve this issue, yet in reality there is a paucity of well-provenanced human fossils from early Upper Paleolithic contexts. One specimen, a right humerus from the site of Vogelherd (Germany), has been argued, based on its size, robusticity, and muscularity, to possibly represent a Neandertal in an Aurignacian context. The morphological affinities of the Vogelherd humerus were explored by univariate and multivariate comparisons of humeral epiphyseal and diaphyseal shape and strength measures relative to humeri of Neandertals and Early Upper Paleolithic (later Aurignacian and Gravettian) modern humans. On the basis of diaphyseal cross-sectional geometry, deltoid tuberosity morphology, and distal epiphyseal morphology, the specimen falls clearly and consistently with European early modern humans and not with Neandertals. Along with the other Vogelherd human remains, the Vogelherd humerus represents an unequivocal association between the Aurignacian and modern human morphology in Europe.

Animals↗

New evidence from Le Moustier 1: computer-assisted reconstruction and morphometry of the skull.

In this study, we present a new computerized reconstruction of the Le Moustier 1 Neanderthal skull and discuss its significance for Neanderthal growth and variability. Because of the precarious state of preservation of the original material, we applied entirely noninvasive methods of fossil reconstruction and morphometry, using a combination of computed tomography, computer graphics, and stereolithography. After electronic restoration, the isolated original pieces were recomposed on the computer screen using external and internal anatomical clues to position the bone fragments and mirror images to complete missing parts. The inferred effects of general compressive deformation that occurred during fossilization were corrected by virtual decompression of the skull. The resulting new reconstruction of the Le Moustier 1 skull shows morphologic features close to the typical Neanderthal adult state. Residual asymmetry of skeletal parts can be traced to in vivo skeletal modification: the left mandibular joint shows signs of a healed condylar fracture, and the anatomy of the occipital region suggests mild plagiocephaly. Using micro-CT analysis, the left incus could be recovered from the matrix filling of the middle ear cavity. Its morphometric dimensions are similar to those of the La Ferrassie III incus. The morphometric characteristics of the inner ear deviate substantially from the condition reported as typical for Neanderthals and fall within the range of modern human variability.

Animals↗

Allosaurus, crocodiles, and birds: evolutionary clues from spiral computed tomography of an endocast.

Because the brain does not usually leave direct evidence of its existence in the fossil record, our view of this structure in extinct species has relied upon inferences drawn from comparisons between parts of the skeleton that do fossilize or with modern-day relatives that survived extinction. However, soft-tissue structure preservation may indeed occasionally occur, particularly in the endocranial space. By applying modern imaging and analysis methods to such natural cranial "endocasts," we can now learn more than ever thought possible about the brains of extinct species. I will discuss one such example in which spiral computed tomography (CT) scanning analysis has been successfully applied to reveal preserved internal structures of a naturally occurring endocranial cast of Allosaurus fragilis, the dominant carnivorous dinosaur of the late Jurassic period. The ability to directly examine the neuroanatomy of an extinct dinosaur, whose modern-day relatives are birds and crocodiles, has exciting implications about Allosaurus' behavior, its adaptive responses to its environment, and its eventual extinction.

Alligators and Crocodiles↗

Developmental mechanisms of vertebrate limb evolution.

Over the past few years, our understanding of the evolution of limbs has been improved by important new discoveries in the fossil record. Additionally, rapid progress has been made in identifying the molecular basis of vertebrate limb development. It is now possible to integrate these two areas of research in order to identify the molecular developmental mechanisms underlying the evolution of paired appendages in vertebrates. After the origin of paired appendages, several vertebrate lineages reduced or eliminated fins and limbs and returned to the limbless condition. Examples include eels, caecilians, snakes, slow worms and several marine mammals. Analyses of fossil and extant vertebrates show that evolution of limblessness frequently occurred together with elongation of the trunk and loss of clear morphological boundaries in the vertebral column. This may be suggestive of a common developmental mechanism linking these two processes. We have addressed this question by analysing python embryonic development at tissue, cellular and molecular levels, and we have identified a developmental mechanism which may account for evolution of limb loss in these animals.

Animals↗

Nipping the Cambrian "explosion" in the bud?

In recent years, two schools of thought have emerged with regard to the Cambrian "explosion". One argues that it was very quick, with phyla tumbling into existence in a virtual geological instant. The other view has a more relaxed temporal perspective. It looks to slow aeons of cryptic metazoan history, which led to a final breakthrough in the Cambrian, not in evolution but of fossilization potential. Yet both views have serious difficulties. Now, in a recent issue of Biological Reviews, Graham Budd and Sören Jensen(1) argue for a third way. In an intriguing blend of functional morphology, the fossil record and cladistic thinking, they suggest that the assembly of metazoan bodyplans took place in a surprisingly straightforward manner.

Animals↗

The limits of palaeontological knowledge: finding the gold among the dross.

Palaeontological interpretation rests on two interwoven sets of comparisons with the modern world. Palaeobiological interpretation relies on the placement of fossils within a phylogenetic and functional framework based primarily on the comparative biology of living organisms. Analogy to currently observable chemical, physical and taphonomic processes enables palaeoenvironmental inferences to be drawn from geological data. In older rocks, comparisons with the modern Earth can become tenuous, limiting palaeontological interpretation. The problem reaches its apogee in Archaean successions, yet pursuit of multiple lines of evidence establishes that complex microbial communities, fuelled by autotrophy and, likely, photoautotrophy, existed 3500 million years ago. Although Archaean palaeontology has to date focused on silicified coastal sediments, improved understanding of Earth's earliest biosphere may depend on the development of alternative environmental and taphonomic analogies. Spring precipitates and hydrothermal metal deposits are promising candidates. Terrestrial organisms may be of limited value in interpreting such fossils as may be found on Mars, although some points of comparison could prove general. Given limited opportunities for exploration, proper choice of environmental analogy is critical. Spring precipitates constitute excellent deposits for addressing questions of biology on another planet.

Archaea↗

Hominid cranial remains from upper Pleistocene deposits at Aduma, Middle Awash, Ethiopia.

The Upper Pleistocene localities of Aduma and Bouri have yielded hominid fossils and extensive Middle Stone Age (MSA) archaeological assemblages. The vertebrate fossils recovered include parts of four hominid crania from Aduma and a complete right parietal from Bouri. Archaeological associations and radiometric techniques suggest an Upper Pleistocene age for these hominids. The more complete cranium from Aduma (ADU-VP-1/3) comprises most of the parietals, the occipital, and part of the frontal. This cranium is compared to late Middle and Upper Pleistocene hominid crania from Africa and the Middle East. The Aduma cranium shows a mosaic of cranial features shared with "premodern" and anatomically modern Homo sapiens. However, the posterior and lateral cranial dimensions, and most of its anatomy, are centered among modern humans and resemble specimens from Omo, Skhul, and Qafzeh. As a result, the Aduma and Bouri Upper Pleistocene hominids are assigned to anatomically modern Homo sapiens.

Animals↗

Brain size and encephalization in early to Mid-Pleistocene Homo.

Important changes in the brain have occurred during the course of human evolution. Both absolute and relative size increases can be documented for species of Homo, culminating in the appearance of modern humans. One species that is particularly well-represented by fossil crania is Homo erectus. The mean capacity for 30 individuals is 973 cm(3). Within this group there is substantial variation, but brain size increases slightly in specimens from later time periods. Other Middle Pleistocene crania differ from those of Homo erectus. Characters of the facial skeleton, vault, and cranial base suggest that fossils from sites such as Arago Cave in France, the Sima de los Huesos in Spain, Bodo in Ethiopia, Broken Hill in Zambia, and perhaps Dali in China belong to the taxon Homo heidelbergensis. Ten of these mid-Quaternary hominins have brains averaging 1,206 cm(3) in volume, and many fall beyond the limits of size predicted for Homo erectus of equivalent age. When orbit height is used to construct an index of relative brain size, it is apparent that the (significant) increase in volume documented for the Middle Pleistocene individuals is not simply a consequence of larger body mass. Encephalization quotient values confirm this finding. These changes in absolute and relative brain size can be taken as further corroborative evidence for a speciation event, in which Homo erectus produced a daughter lineage. It is probable that Homo heidelbergensis originated in Africa or western Eurasia and then ranged widely across the Old World. Archaeological traces indicate that these populations differed in their technology and behavior from earlier hominins.

Animals↗

Genetics and the evolution of primate enamel thickness: a baboon model.

The thickness of mammalian tooth enamel plays a prominent role in paleontology because it correlates with diet, and thicker enamel protects against tooth breakage and wear. Hominid evolutionary studies have stressed the importance of this character for over 30 years, from the identification of "Ramapithecus" as an early Miocene hominid, to the recent discovery that the earliest hominids display molar enamel intermediate in thickness between extant chimpanzees and Australopithecus. Enamel thickness remains largely unexplored for nonhominoid primate fossils, though there is significant variation across modern species. Despite the importance of enamel thickness variation to primate evolution, the mechanisms underlying variation in this trait have not yet been elucidated. We report here on the first quantitative genetic analysis of primate enamel thickness, an analysis based on 506 pedigreed baboons from a captive breeding colony. Computed tomography analysis of 44 Papio mandibular molars shows a zone of sufficiently uniform enamel thickness on the lateral surface of the protoconid. With this knowledge, we developed a caliper metric measurement protocol for use on baboon molars worn to within this zone, enabling the collection of a data set large enough for genetic analyses. Quantitative genetic analyses show that a significant portion of the phenotypic variance in enamel thickness is due to the additive effects of genes and is independent of sex and tooth size. Our models predict that enamel thickness could rapidly track dietary adaptive shifts through geological time, thus increasing the potential for homoplasy in this character. These results have implications for analyses of hominoid enamel thickness variation, and provide a foundation from which to explore the evolution of this phenotype in the papionin fossil record.

Animals↗

Alternative dental measurements: proposals and relationships with other measurements.

Most archaeological and fossil teeth are heavily worn, and this greatly limits the usefulness of tooth crown diameter measurements, as they are usually defined at the widest points of the crown. There are alternatives, particularly measurements at the cervix of the tooth, where the crown joints the root, and measurements along a diagonal axis in molars, that are much less affected by wear. These would allow a wider range of specimens to be included, e.g., in the study of dental reduction in Upper Palaeolithic and Mesolithic Homo sapiens. In addition, they would allow the little-worn teeth of children to be compared directly with well-worn teeth in adults. These alternatives, however, have been little used, and as yet there have not been any studies of the repeatability with which they can be measured, or of the extent to which they are related to the more usual crown diameters. The present study is based on a group of unworn teeth, where direct comparisons could be made between the alternative measurements, which are not much affected by wear, with the usual crown diameters, which are very much affected. In an interobserver-error study of this material, cervical and diagonal measurements could be recorded as reliably as the usual crown diameters. The buccolingual cervical measurement was strongly correlated with the normal bucclingual crown diameter in all teeth, whereas the mesiodistal cervical measurement was highly correlated with the normal mesiodistal crown diameter in incisors and canines, but less so in premolars and molars. The molar diagonal measurements showed high correlations with all other measurements. Crown areas (robustness index) calculated from the usual diameters were strongly correlated with crown areas calculated from cervical measurements, and crown areas calculated from molar diagonals were strongly correlated with both other areas. Despite the long usage of the more usual maximum crown diameters, the alternative dental measurements could be measured just as reliably, could record similar information about tooth crown size, and would be better measures for the worn dentitions seen in archaeological and fossil material.

Fossils↗

Craniodental variation in Paranthropus boisei: a developmental and functional perspective.

What levels and patterns of craniodental variation among a fossil hypodigm are necessary to reject the null hypothesis that only a single species is sampled? We suggest how developmental and functional criteria can be used to predict where in the skeleton of fossil hominins we should expect more, or less, within-species variation. We present and test three hypotheses about the factors contributing to craniodental variation in extant primate taxa, and then apply these results to the interpretation of the P. boisei hypodigm. Within the comparative samples of extant Homo, Pan, Gorilla, Pongo, and Colobus, variables from the cranial base, neurocranium, and face that are not subject to high magnitudes of strain have consistently lower levels of intraspecific variation than variables from regions of the face subject to high levels of strain. Dental size variables are intermediate in terms of their reliability. P. boisei is found to have a low degree of variability relative to extant primates for variables shown to be generally useful for testing taxonomic hypotheses. Contrary to the claims of Suwa et al. ([1997] Nature 389:489-492), the recently discovered material from Konso falls within the range of variation of the "pre-Konso" hypodigm of P. boisei for available conventional metrical variables. Those aspects of the Konso material that appear to extend the range of the P. boisei hypodigm involve regions of the skull predicted to be prone to high levels of within-species variation. The approach used in this study focuses on craniodental data, but it is applicable to other regions of the skeleton.

Animals↗

Aspects of the middle and external ear of early South African hominids.

A brief description of the exposed middle ear from two Swartkrans specimens demonstrates the feasibility of examining the inner auditory structures of existing fossil hominid material. These Swartkrans fossils provide the first clear view of the early hominid middle ear, and their structures appear to be morphologically distinct. The bony anatomy of the external ear of Australopithecus robustus is found to be unique among the South African hominids.

Animals↗

Engis: preparation damage, not ancient cutmarks.

Scratches found on the Engis 2 cranium have been described as perimortem and interpreted as intentional scalping marks by Russell and LeMort (Am. J. Phys. Anthropol. 69:317-323, 1986). These marks are described and compared to damage on other fossil hominids. The Engis marks have been misinterpreted. These marks are sandpaper striae formed during restoration of the vault, moulding striae formed when mold part lines were incised into the fossil and profiling striae formed when craniograms were made with sharp steel instrument tips. None of them have anything to do with prehistoric behavior.

Animals↗