Early Acheulean technology in the Rietputs Formation, South Africa, dated with cosmogenic nuclides

An absolute dating technique based on the build-up and decay of ²⁶Al and ¹⁰Be in the mineral quartz provides crucial evidence regarding early Acheulean hominid distribution in South Africa. Cosmogenic nuclide burial dating of an ancient alluvial deposit of the Vaal River (Rietputs Formation) in the western interior of South Africa shows that coarse gravel and sand aggradation there occurred ca 1.57 ± 0.22 Ma, with individual ages of samples ranging from 1.89 ± 0.19 to 1.34 ± 0.22 Ma. This was followed by aggradation of laminated and cross-bedded fine alluvium at ca 1.26 ± 0.10 Ma. The Rietputs Formation provides an ideal situation for the use of the cosmogenic nuclide burial dating method, as samples could be obtained from deep mining pits at depths ranging from 7 to 16 meters. Individual dates provide only a minimum age for the stone tool technology preserved within the deposits. Each assemblage represents a time averaged collection. Bifacial tools distributed throughout the coarse gravel and sand unit can be assigned to an early phase of the Acheulean. This is the first absolute radiometric dated evidence for early Acheulean artefacts in South Africa that have been found outside of the early hominid sites of the Gauteng Province. These absolute dates also indicate that handaxe-using hominids inhabited southern Africa as early as their counterparts in East Africa. The simultaneous appearance of the Acheulean in different parts of the continent implies relatively rapid technology development and the widespread use of large cutting tools in the African continent by ca 1.6 Ma.     

Cut marks on the Bodo cranium: a case of prehistoric defleshing

Cut marks were discovered on the Middle Pleistocene Bodo cranium from Ethiopia. The cut marks most closely resemble experimental damage caused by the application of stone tools to fresh bone. This discovery constitutes the earliest solid evidence for intentional defleshing of a human ancestor and offers new research avenues for the investigation of early hominid mortuary practices.     

First occurrence of early Homo in the Nachukui Formation (West Turkana, Kenya) at 2.3-2.4 Myr

Cognitive abilities and techno-economic behaviours of hominids in the time period between 2.6-2.3 Myr have become increasingly well-documented. This time period corresponds to the oldest evidence for stone tools at Gona (Kada Gona, West Gona, EG 10-12, OGS 6-7), Hadar (AL 666), lower Omo valley (Ftji1, 2 & 5, Omo 57, Omo 123) in Ethiopia, and West Turkana (Lokalalei sites -LA1 & LA2C-) in Kenya. In 2002 a new palaeoanthropological site (LA1alpha), 100 meters south of the LA1 archaeological site, produced a first right lower molar of a juvenile hominid (KNM-WT 42718). The relative small size of the crown, its marked MD elongation and BL reduction, the relative position of the cusps, the lack of a C6 and the mild expression of a protostylid, reinforced by metrical analyses, demonstrate the distinctiveness of this tooth compared with Australopithecus afarensis, A. anamensis, A. africanus and Paranthropus boisei, and its similarity to early Homo. The LA1alpha site lies 2.2 m above the Ekalalei Tuff which is slightly younger than Tuff F dated to 2.34+/-0.04 Myr. This juvenile specimen represents the oldest occurrence of the genus Homo in West Turkana.     

Modeling the formation of Early Stone Age artifact concentrations

The nature of stone artifact concentrations at early Plio-Pleistocene sites in East Africa is evaluated with regard to hominid transport behaviors responsible for their formation. These archaeological occurrences indicate ranging behaviors involving deliberate and repeated transport of flaked stone artifacts. The stone transported to archaeological sites within the time range of Homo habilis indicates planned transport of tools or material for tool manufacture to an extent far beyond transport behaviors reported among living apes, even stone hammer-using chimpanzees. Analysis of technological evidence in a lithic assemblage at a Plio-Pleistocene site at Koobi Fora (c. 1·5 ya) indicates on-site manufacturing activities and transport of flaked stone both to and from the site locale. Possible explanations for transport of stone artifacts are discussed in view of hominid strategies of environmental exploitation and resource utilization. A model is proposed for planned, habitual transport of artifacts by hominids positively correlated with distance of planned foraging range. In this model, larger-scale sites tended to develop at locales favorably located near abundant resources, where stone imports were high but export was relaxed due to the proximity of resources to be processed.     

Les sites archéologiques plio-pléistocènes de la formation de Nachukui, Ouest-Turkana, Kenya : bilan synthétique 1997–2001

Plio-Pleistocene archaeological sites in the Nachukui Formation, West Turkana, Kenya: synthetic results 1997–2001. Stretched along the western side of the Turkana Basin, the Nachukui Formation preserves a large number of Plio-Pleistocene archaeological sites. Research carried out by the WTAP documents hominid behavioral evolution and technical diversity, through a time period ranging from 2.35 Myr to 0.7 Myr and within a relatively precise chronostratigraphic and paleoenvironmental frame. This work is based on comprehensive excavations conducted between 1997 and 2001 at the Late Pliocene and Early Pleistocene site complexes of the formation. To cite this article: H. Roche et al., C.R. Palevol 2 (2003).     

Evolution of Hominid Femur and Tibia: A morphometric approach to the evolutionary research in anthropology

Some theoretical and methodological morphometrical approaches in evolutionary anthropology and paleoanthropology are reviewed in this study. It is shown which are the contemporary possibilities of sophisticated biometrical and biostatistical methods and the role of the morphometrical approach. A new approach, experimental morphometrics, is presented, reflecting recent trends in evolutionary morphology as well as sophisticated biostatistical methods. The approach emphasizes the complex inter-related approach to the data processing and a double nature of morphometric data, i.e. biological and biostatistical one. The practical use of experimental morphometry is given for the two examples of analyses of the evolution of the hominoid and hominid femur and tibia. The hypothesis on a two stage restructuring of morphology of the hominid femur and tibia is supported by experimental results. Two different steps during this restructuring could be recognized: 1) Structural remodelling typical for the origin of hominids and australopithecine evolution, and 2) proportional remodelling of lower limb long bones which is connected with the Australopithecus/Homo transition (i.e. mainly Homo habilis stage). The results confirm the increasing trend of bipedal adaptations on the early hominid lower limb skeleton. Analysis of microevolutionary trends on the Homo sapiens femur and tibia indicates at least three different morphological patterns, Paleolithic, Neolithic and Recent, with numerous specific features in morphology and proportions. Neanderthal morphology is very derived. Upper Paleolithic/Mesolithic/Neolithic transition has a key character for the understanding of post-Paleolithic morphology. A very high sexual dimorphism of the femur and tibia has been demonstrated for Upper Paleolithic and Neolithic populations. Presented at the Foundation of Different Approaches to the Study of Human Evolution edited by B. Sigmon & V.V. Leonovicova-Liblice, September 1–3, 1989     

A Pelican Tarsometatarsus (Aves: Pelecanidae) from the Latest Pliocene Siwaliks of India

We report a new fossil specimen of a pelican from the Tatrot Formation of the Siwalik Hills, India. It likely represents Pelecanus sivalensis Davies, 1880, the smaller of the two previously published species from the Siwalik Group stratigraphic sequence. This complete tarsometatarsus is the first fossil bone of a pelican collected in India for over 100 years. It is from the latest Pliocene (∼2.6 Ma), and is the youngest pelican fossil from the region. The new specimen exhibits a derived distoplantar ‘slant’ to the plantar margin of the medial crest of the hypotarsus, and a combination of features related to the morphology of the hypotarsus, the distal foramen, trochleae, and overall size that allow further differentiation from known tarsometatarsi of fossil and extant pelicans, including the three species of extant pelicans that occur in India (Pelecanus crispus, P. onocrotalus, and P. philippensis). It is of appropriate size for Pelecanus sivalensis, which to date has been known only by fragments of other skeletal elements of the wing, leg, and shoulder girdle. Thus, the observation that this tarsometatarsus is morphologically distinct from those of known pelicans provides further support for the distinctiveness of at least one extinct species of pelican from the Siwalik Group sediments. While the morphology of the tarsometatarsus allows for separation from other taxa known from tarsometatarsi, we found no clear shared derived states to place this taxon with any confidence in a phylogenetic context relative to any other pelican species, or even determine if it is part of the crown group of Pelecanidae. However, published molecular data are consistent with an origin of the crown clade prior to the Pleistocene, suggesting (along with one morphological character) the possibility that this species belongs to the Old World clade of pelican species.     

New small mammals from the Hasnot–Tatrot area of the Potwar Plateau,northern Pakistan

New rodent and lagomorph fossils from the Hasnot–Tatrot area of northern Pakistan are presented here to complement knowledge of stratigraphic ranges and morphology of key late Neogene Siwalik taxa. Most of the material is from two sites near the village of Bhandar in strata of the late Miocene age Dhok Pathan Formation; one specimen comes from the Pliocene Tatrot beds. We apply previously established magnetostratigraphy to date the fossils, the Bhandar sites dating to 6.6–6.7 Ma. In describing the fossils, we emphasize new morphological information represented by the material. As surface finds, these fossils represent relatively large body size species: three bamboo rat relatives, a porcupine, and a rabbit. The bamboo rats (Rhizomyinae) are an endemic group, and both the porcupine (Hystrix) and the rabbit (Alilepus) represent late Miocene immigrants into the Indian subcontinent.     

Length estimate for KNM-ER 736, a hominid femur from the lower pleistocene of East Africa

Our knowledge concerning stature in earlyHomo is scanty. In this paper, based on comparison with the fossil femur KNM-ER 999, an estimate of 482 mm femur length is derived for KNM-ER 736, the latter dating from the Lower Pleistocene. From comparison with other fossil and modern femora, KNM-ER 736 appears to be the longest hominid femur so far recovered from a site of Early Pleistocene age. Moreover, the estimated femur length is higher than the published mean values of most modern populations. Provided that trunk and head proportions were not radically different from modernH. sapiens, the finding would suggest that a stature similar to that of modern man was already reached by East AfricanHomo as early as about 1.6 Myr before present.     

Nouvelles découvertes de dents d’hominidés dans le membre Kaitio de la formation de Nachukui (1,65–1,9 Ma), Ouest du lac Turkana (Kenya)

New hominid teeth from the Kaitio member (1.65–1.9 Myr) in West Turkana (Kenya). New hominid teeth have been recovered from the archaeological sites of Kokiselei 1 and Naiyena Engol 1. These two sites are located in the west side of the Turkana Basin and belong to the Kaitio member of the Nachukui Formation. They are dated between 1.65-1.79 and 1.7-1.8 Myr respectively. The four teeth (left maxillary canine and first molar, right maxillary third molar and left mandibular third molar) discovered in Kokiselei 1 are attributed to Australopithecus boisei. The right mandibular first premolar found in Naiyena Engol 1 is referred to Homo sp. aff. ergaster. To cite this article: S. Prat et al., C. R. Palevol 2 (2003).     

Soanian lithic occurrences and raw material exploitation in the Siwalik Frontal Zone, northern India: a geoarchaeological perspective

In the Himalayan foothills of northern India, evidence of widespread hominin occupation since at least the late Middle Pleistocene has been known since the early 20th century and indicates varied patterns of land-use and intraregional mobility. This lithic evidence primarily belongs to the Soanian industry, representing some of the highest concentrations of Paleolithic assemblages in the Old World based exclusively on pebble and cobble clasts. This body of evidence also signifies interregional dispersal from peninsular India or northern Pakistan, leading to environmental preferences that spread quickly through hominin populations in the region within a relatively short timespan. While rich in its technological repertoire, the Soanian industry is poorly- understood regarding site selection and raw material exploitation over time. Recent efforts demonstrate that Soanian sites on Siwalik frontal slopes between two major rivers vary considerably in their artifact quantities regardless of abundant raw material sources found across the landscape. Most of the assemblages suggest raw material transport distances of three kilometers or less from the localized sources. Geoarchaeological investigations at the richest known Soanian site, Toka, reveal dynamic evidence of pre- and postdepositional site formation including the exploitation of quartzite pebbles and cobbles by Pleistocene hominins from terrace and streambed contexts within a 1 km2 radius. Some field observations also disprove claims made by previous workers, of artifacts eroding out of late Pliocene exposures of the Upper Siwalik Tatrot Formation around Toka.     

Lithic microwear analysis in archeology

Stone tools are the most durable and ubiquitous residue of prehistoric hominid activity. For this reason, archeologists attempt to learn as much as possible about hominid behavior from the analysis of lithic artifacts. Lithic microwear analysis reconstructs aspects of stone-tool use from patterned variation in the traces of microscopic wear on those tools. The analysis of lithic microwear traces has increased our understanding of how stone tools were used in contexts ranging from the early Pleistocene to the ethnographic present.     

Antelopes (Mammalia,Ruminantia, Bovidae) from the Upper Siwaliks of Tatrot,Pakistan, with description of a new species

New antelope material from the Tatrot type locality of the Tatrot Formation, Upper Siwaliks, northern Pakistan, allows us to recognize four antelopes, i.e., Antilope intermedia sp. nov. Khan and Akhtar, Antilope subtorta Pilgrim, 1937, Antilope cervicapra (Linnaeus, 1758), and Kobus porrecticornis (Lydekker, 1878). Comparisons with A. cervicapra and A. subtorta suggest that A. intermedia sp. nov. is closely related to the black buck A. cervicapra. The new material from Tatrot confirms the idea that antelopes in the Upper Pliocene Tatrot Formation were rather diverse.     

Evolution of femur and tibia in higher primates: Adaptive morphological patterns and phylogenetic diversity

Seventy six metrical traits measured on the femur and tibia of three higher primate groups —Ceboidea, Cercopithecoidea, Hominoidea have been processed by various univariate and multivariate statistical methods to survey the process of evolution of the morphology of the femur and tibia in higher primates. Intragroup and intergroup variability, similarity and differences as well as various aspects of scaling and sexual dimorphism have been analyzed to study adaptive trends and phylogenetic diversity in higher primates, in individual superfamilies and to explore the adaptive morphological pattern of early hominids and basic differences between hominids and pongids. Two basic morphotypes of the femur and tibia in higher primates have been determined. They are (1) advanced hominoid morphotype (hominids and pongids) and (2) ancestral higher primate morphotype (platyrrhine and cattarrhine monkeys, early hominoids, and hylobatids). Cebid lower limb bones are adapted to arboreal quadrupedalism with antipronograde features while femur and tibia of cercopithecid monkeys are basically adapted to the semi-arboreal locomotion. Early hominoids (Proconsul) and hylobatids are morphologically different from pongids; some features are close toAteles or other monkey species. Pongids and hominids are taken as one major morphological group with different scaling and some functional and morphological similarities. Numerous analogous features were described on the lower limb skeleton ofPan andPongo showing analogous ecological parameters in their evolution. Major morphological and biomechanical trends are analyzed. It is argued that early advanced hominoid morphology is ancestral both to the pongids and to early hominids. The progressive morphological trend in early hominids has been found fromA. afarensis with ancestral hominid morphology, toH. habilis with an elongated femur and structural features similar to advanced hominids. A detailed phylogenetic analysis of higher primate femur and tibia is also presented.     

Taphonomic perspectives on hominid site use and foraging strategies during Bed II times at Olduvai Gorge, Tanzania

The faunal assemblages excavated by Mary Leakey in Bed II of Olduvai Gorge, Tanzania, have, like the more well-known Bed I assemblages, traditionally been interpreted as the result of hominid butchering activities in the lake margin and riverine settings of the paleo-Olduvai Basin. A reexamination of all of Leakey's Bed I sites has shown that hominids played little or no role in the formation of all but one of those faunal assemblages, a finding that prompted the reanalysis of the Bed II sites presented here. We expand upon a previous taphonomic study that provided systematic data for HWK East Levels 1–2, MNK Main, and BK. In addition to these assemblages, we provide data on HWK East Levels 3–5, FC West, TK, and SHK. Our data contradict previous interpretations of MNK Main as a hominid accumulation but uphold the contention that BK represents a primarily hominid accumulation reflecting early access to carcasses. The small and poorly preserved assemblages from FC West and TK are difficult to link unambiguously to either hominids or carnivores. Site MNK Main and HWK East Levels 3–5 appear to be death arenas where carcasses accumulated via natural deaths and/or serial predation. Site SHK is severely biased by selective retention and therefore little can be said of its formational history. Nevertheless, no hominid modifications were documented in this assemblage. Comparisons with other Olduvai sites indicate a more conspicuous hyena taphonomic signal during Bed II times than Bed I times, which appears to mirror the changing configuration of the large carnivore guild. These findings also beg the question of what activities were being carried out by hominids with the stone tools discarded at these sites. Although it seems clear that hominids were utilizing stone tools to carry out subsistence activities unrelated to carcass butchery, more excavation and techniques such as phytolith analysis should be employed to explore alternative explanations.     

Femur length,body mass,and stature estimates of <Emphasis Type="Italic">Orrorin tugenensis</Emphasis>, a 6 Ma hominid from Kenya

To understand the palaeobiology of extinct hominids it is useful to estimate their body mass and stature. Although many species of early hominid are poorly preserved, it is occasionally possible to calculate these characteristics by comparison with different extant groups, by use of regression analysis. Calculated body masses and stature determined using these models can then be compared. This approach has been applied to 6 Ma hominid femoral remains from the Tugen Hills, Kenya, attributed to Orrorin tugenensis. It is suggested that the best-preserved young adult individual probably weighed approximately 35–50 kg. Another fragmentary femur results in larger estimates of body mass, indicative of individual variation. The length of the femur of the young adult individual was estimated, by using anthropoid-based regression, to be a minimum of 298 mm. Because whole-femur proportions for Orrorin are unknown, this prediction is conservative and should be revised when additional specimens become available. When this predicted value was used for regression analysis of bonobos and humans it was estimated to be 1.1–1.2 m tall. This value should, however, be viewed as a lower limit.     

Two new“Meganthropus” mandibles from Java

There are now eleven known mandibular pieces from the Lower and Middle Pleistocene of Java, all but one being from the Sangiran site. All of these have been assigned toHomo erectus by most authorities, while others have suggested as many as four different hominoid taxa. Two of the mandibles, Sangiran 33 (Mandible H) and“Meganthropus”D (no Sangiran number yet assigned), are described here for the first time. The two new mandibles come from the Upper Pucangan Formation and date approximately 1.2–1.4 Myr. They are morphologically compatible with other“Meganthropus” mandibles described from Java. Despite attempts by numerous authorities to place all the Sangiran hominid mandibles in the species,H. erectus, the range of variation in metric and nonmetric features of the“Meganthropus” hominids is clearly beyond the know variation found inH. erectus. “Meganthropus” could represent a speciation from the well-knownH. erectus.     

Behavioral inferences from Early Stone artifact assemblages: an experimental model

A methodological approach for assessing the nature and palaeographic distribution of early stone artifact assemblages is presented, modeled after an approach originally used for faunal analysis. By combining experimental replicative studies with careful analysis of Palaeolithic archaeological occurrences, it is potentially possible to reconstruct entire technological systems and to assess what stages of lithic reduction may be preferentially represented at high density artifact concentrations that we normally call archaeological “sites”, and what stages of reduction are found in lower density (“off site”) scatters.Based upon the results of this approach, alternative explanations for certain stages of lithic reduction being preferentially represented at a number of Plio-Pleistocene archaeological sites at Koobi Fora, Kenya are considered and evaluated with regard to early hominid organizational patterns. It appears that hominid stone technology was a relatively complex system by 1·9 to 1·4 million years B.P., involving significant transport and carrying of stone artifacts representing various stages of reduction and suggesting more foresight and planning than observed among extant nonhuman primates.The application of this approach to other Palaeolithic occurrences should enable anthropologists to obtain a better understanding of the organizational patterns of tool-making hominids throughout the course of human evolution.     

On the relative frequencies of hominid maxillary and mandibular teeth and jaws as taphonomic indicators

In southern African samples of early hominid remains, maxillary and mandibular teeth (deciduous-plus-permanent) have a virtually equal chance of accumulating in the dolomitic limestone cave deposits, of being preserved therein and recovered therefrom. Thus, of 1066 fossil teeth ofAustralopithecus spp. plusHomo habilis, 51.9 per cent are maxillary and 48.1 per cent mandibular. On the other hand, the East African sample of 847 early hominid, deciduous-plus-permanent teeth, departs more strikingly from a 1∶1 ratio: it comprises 41.0 per cent maxillary and 59.0 per cent mandibular teeth. It is inferred that mandibular teeth have a somewhat better chance of accumulating and being preserved in, and being recovered from, the open, fluvial, lacustrine and deltaic sedimentary environments of the East African sites. The dental proportions are approximately matched by the proportions of jaws. For example, the maxilla: mandible proportions at Koobi fora in northern Kenya are 33.0∶67.0 for teeth and 21.6∶78.4 for jaws. In other words, the preponderance in favour of mandibular remains is somewhat more marked in the case of jaws than of teeth, this distinction doubtless reflecting the more fragile bony structure of the maxilla and the sturdier construction of the mandible. This first study known to the author of the differential distribution of maxillary and mandibular teeth of the Plio-Pleistocene hominids leads the author to hypothesize that, where environmental conditions at the place and time of the death of the hominids have been non-destructive, non-dispersive, relatively mild and protective, maxillae and mandibles may be expected to have been conserved and recovered in approximately equal proportions—and likewise of maxillary and mandibular teeth. On the other hand, the more brutal and destructive the sedimentary environment and other taphonomic influences have been, at the place and time when the hominid individuals died, the more likely it is that the maxillary and mandibular remains of jaws and teeth will deviate from equality of proportions, generally at the expense of the maxillae and upper teeth. Hence, it is proposed that the upper jaw/low jaw ratio (Mx/Mn jaw ratio) and the maxillary teeth/mandibular teeth ratio (Mx/Mn dental ratio) may serve as two useful new gauges of the rigour of palaeo-ecological and taphonomic conditions.     

The neural crest and neural crest cells: discovery and significance for theories of embryonic organization

The neural crest has long fascinated developmental biologists, and, increasingly over the past decades, evolutionary and evolutionary developmental biologists. The neural crest is the name given to the fold of ectoderm at the junction between neural and epidermal ectoderm in neurula-stage vertebrate embryos. In this sense, the neural crest is a morphological term akin to head fold or limb bud. This region of the dorsal neural tube consists of neural crest cells, a special population(s) of cell, that give rise to an astonishing number of cell types and to an equally astonishing number of tissues and organs. Neural crest cell contributions may be direct — providing cells — or indirect — providing a necessary, often inductive, environment in which other cells develop. The enormous range of cell types produced provides an important source of evidence of the neural crest as a germ layer, bringing the number of germ layers to four — ectoderm, endoderm, mesoderm, and neural crest. In this paper I provide a brief overview of the major phases of investigation into the neural crest and the major players involved, discuss how the origin of the neural crest relates to the origin of the nervous system in vertebrate embryos, discuss the impact on the germ-layer theory of the discovery of the neural crest and of secondary neurulation, and present evidence of the neural crest as the fourth germ layer. A companion paper (Hall, Evol. Biol. 2008) deals with the evolutionary origins of the neural crest and neural crest cells.