Hominid fossil sample from Kanjera,Kenya: Description,provenance, and implications of new and earlier discoveries

Anatomically modern hominids were first collected from Kanjera, Kenya, by L.S.B. Leakey in the 1930s. Their apparent association with an archaic fauna was quickly challenged, throwing their age into doubt. Further unpublished hominid fragments were collected in 1974, 1975, 1981, and 1987. We review the context and morphology of the entire hominid sample. A minimum number of five individuals is represented by both cranial and postcranial elements. Several individuals have thickened cranial vaults, a characteristic originally thought to reflect their great antiquity. Vault thickening resulted from diploic expansion and may have been a response to acquired or inherited anemia. The entire hominid sample postdates the Kanjera Formation, deposited from the early into the middle Pleistocene. Most of the sample was derived from the black cotton soil capping the stratigraphic column. The morphology and context of the Kanjera hominids is consistent with human skeletal remains from nearby Holocene sites. Hominid 3 was probably an intrusive burial into an early Pleistocene bed. © 1995 Wiley-Liss, Inc.     

Récentes découvertes de restes d'Hominidés fossiles en Chine du Sud

After the discovery of the controversial hominids and artefacts at the Longgupo site, three Early Pleistocene sites in the southern valleys of the Changjiang (Yangtze) were excavated from 1998 to 2000 in order to test the hypothesis that the hominids exist in China before 2 Ma. Three cheek teeth of Meganthropus palaeojavanicus, more than six hundreds pieces of artefacts of Mode 1 technology and thousands pieces of mammal fossils of Villafranchian age were unearthed in situ at the Longgudong Site in Hubei Province. More than 120 pieces of artefacts of Mode 1 technology and seven thousands pieces of mammal fossils of Villafranchian age were unearthed in situ at the Renzidong Site in Anhui Province. Although no new materials of hominids and artefacts were unearthed at the Yuanmou Man Site at Danawu in Yunnan Province, new materials of mammalian fauna confirm the horizon yielding the Yuanmou Man fossils is of the Early Pleistocene. These new discoveries imply that the appearance of hominids in China is very likely before 2 Ma. If the new report of 3 Ma artefact from Yuxian of the Nihewan Basin in northern China can be confirmed, it will be a strong support for the Continuity Theory.     

Uplift of the roof of africa and its bearing on the evolution of mankind

Evidence concerning the geomorphological evolution of the Western Rift Valley, sedimentation within the valley and comparison of the fossil mammalian faunas of Western Uganda and East Africa indicate that the mountain ranges which now flank the Western Rift were uplifted in three or more stages beginning during the upper Miocene and that they reached climatically important altitudes during the upper Pliocene, at which time they began to modify regional climatic patterns in East Africa. Their main effect was the xerification of conditions over much of the region east of the mountains. The regional climatic effects due to the mountain ranges were themselves modified by global climatic changes related to the onset of the Glacial Period, the two phenomena combining to yield the Present day climatic regime of East Africa. As the climate changed, so did the flora and fauna. Faunal response was of three main kinds: a) dispersal into East Africa of pre-existing forms already adapted to more xeric conditions (many bovids, some cercopithecids), b) autochthonous evolution of forms adapted to mesic environments into forms adapted to more xeric conditions (suids, elephantids, some bovids, hominids), c) displacement of species ranges of those lineages unable to adapt to changing conditions (i.e. local extinctions) (Anancus, Brachypotherium). Autochthonous evolvers, including hominids, adopted two main strategies reflected in their hard anatomy: a) dietary shift (suids, proboscideans, bovids and later Pliocene hominids) and b) locomotor changes (early Pliocene hominids).     

Cutmarked bones from Pliocene archaeological sites at Gona, Afar, Ethiopia: implications for the function of the world's oldest stone tools

Newly recorded archaeological sites at Gona (Afar, Ethiopia) preserve both stone tools and faunal remains. These sites have also yielded the largest sample of cutmarked bones known from the time interval 2.58-2.1 million years ago (Ma). Most of the cutmarks on the Gona fauna possess obvious macroscopic (e.g., deep V-shaped cross-sections) and microscopic (e.g., internal microstriations, Herzian cones, shoulder effects) features that allow us to identify them confidently as instances of stone tool-imparted damage caused by hominid butchery. In addition, preliminary observations of the anatomical placement of cutmarks on several of the recovered bone specimens suggest that Gona hominids may have eviscerated carcasses and defleshed the fully muscled upper and intermediate limb bones of ungulates--activities that further suggest that Late Pliocene hominids may have gained early access to large mammal carcasses. These observations support the hypothesis that the earliest stone artifacts functioned primarily as butchery tools and also imply that hunting and/or aggressive scavenging of large ungulate carcasses may have been part of the behavioral repertoire of hominids by c. 2.5 Ma, although a larger sample of cutmarked bone specimens is necessary to support the latter inference.     

Zooarchaeological and taphonomic analysis of the Die Kelders Cave 1 layers 10 and 11 Middle Stone Age larger mammal fauna

Middle Stone Age (MSA) and Middle Paleolithic (MP) faunal assemblages have gained widespread attention due to their relevance to the debate over the modernity of hominid behavior during the MSA/MP. A recent critique of the scavenging argument for MSA/MP behavior drew on a summary presentation of the skeletal abundance and surface modification data from Die Kelders Cave 1 Layer 10 (Marean, 1998). This paper provides a more complete presentation of those data, adds the smaller Layer 11 sample, and provides a detailed analysis of the taphonomic history of both samples.Bone fragment density is higher in Layer 10 than in Layer 11. Bone densities vary horizontally as well, with Layer 10 showing greater deposition in the exposed areas of the cave. An analysis of long bone breakage patterns indicates that non-nutritive breakage on the Layers 10 and 11 samples was present but not intense. Size 1 mammals were predominantly accumulated by owls and/or other large raptors, not hominids, in Layer 10. Hominids were the predominant accumulator of Sizes 2-4 mammals in Layers 10 and 11 as indicated by the frequency of hammer-stone percussion marks and carnivore toothmarks. After discard by hominids, a significant portion of these remains were discovered and scavenged by carnivores. Overall, the larger mammal fauna of Layer 10 is dominated by Sizes 3 and 4 bovids, mostly young and adult eland, and thus hominids were focusing on the high-ranked prey items. Shaft portions of long bones, the portions with the most flesh, have the highest frequencies of cutmarks. A comparison of the Layers 10 and 11 cutmark frequencies to Selvaggio's (1998) scavenging model shows that the frequencies are significantly outside the range of variation documented in Selavaggio's scavenging sample.     

The environmental background of hominid emergence and the appearance of the genusHomo

The human biologist usually considers ecology of recent humanity. This essay explores the question of whether the human biologist specialising in the ecology of living peoples has anything to learn from the palaeo-anthropologist, studying ancient hominids andtheir adaptive mechanisms over a deep time dimension. Since the Hominidae are under discussion, the definition of the hominids is reviewed. Historically, three phases are recognised. A rethinking of the classification of the hominoids has become necessary for the old and classical systematics, which divided this superfamily into the Hominidae and the Pongidae, is now outmoded. Since no consensus on such a re-classification has yet been reached, the author adheres to the classical system for the time being. The Hominidae emerged between about 8 and 5 million years ago. At that time, Africa was subject to major cooling and aridification and considerable changes in the flora and fauna were occurring. Wet forests were retreating, savanna was spreading and the animals of Africa were undergoing many changes, partly by faunal interchange with Asia following the drying up of the Mediterranean, and partly by autochthonous evolution among the pre-existing species of the continent. The Hominidae could well have emerged from the striking environmental modifications of this late Miocene phase. Critical changes occurred in hominid evolution between 3 and 2 million years before the present. The pre-existing speciesAustralopithecus africanus acquired the form of a postulated derivedA. africanus; the hominid lineage underwent cladogenetic splitting into robust and hyper-robust australopithecines and the genusHomo; Homo babilis appeared; stone tools are first found in the archaeological record; spoken language seems to have been acquired. These sensational events, within the space of one million years, took place against the background of conspicuous changes in the climate and physical geography of Africa, the flora and non-hominid fauna. Mankind became increasingly dependent upon stone culture. Hence a new element was added to the range of modes of adjustment, an element which must have greatly increased the ecological flexibility of the hominids. From the end of the Pliocene era onwards, culture should be seen as a constituent of man's environment and, at the same time, a highly advantageous component of human adaptational processes. In later and recent mankind, it may be difficult to extricate the respective roles of biological, social and physical factors, on the one hand, and cultural aspects on the other, as mechanisms and facilitators of adaptation to diverse econiches.     

Paleoanthropology of the Kibish Formation, southern Ethiopia: Introduction

Cranial and skeletal remains of modern humans, Homo sapiens, were discovered in the Kibish Formation in 1967 by a team from the Kenya National Museums directed by Richard Leakey. Omo I, from Kamoya's Hominid Site (KHS), consists of much of a skeleton, including most of the cranial vault, parts of the face and mandible, and many postcranial elements. Omo II, from Paul's Hominid Site (PHS), is a virtually complete calvaria. Only a limited fauna and a few stone artifacts attributed to the Middle Stone Age were recovered in conjunction with the fossil hominids. The available dating techniques suggested a very early age, over 100 ka, for Member I, from which the Omo I and Omo II fossils were recovered. However, in subsequent decades, the reliability of the dates and the provenance of the Kibish hominids were repeatedly questioned. The papers in this volume provide a detailed stratigraphic analysis of the Kibish Formation and a series of new radiometric dates that indicate an age of 196 +/- 2 ka for Member I and 104 +/- 1 for Member III, confirming the antiquity of the lower parts of the Kibish Formation and, in turn, the fossils from Member I. Studies of the postcranial remains of Omo I indicate an overall modern human morphology with a number of primitive features. Studies of an extensive lithic record from Members I and III indicate a Middle Stone Age technology comparable to assemblages of similar age elsewhere in Ethiopia. Studies of the mammalian, avian, and fish faunas indicate overall similarities to those found in the region today, with a few distinctive differences.     

Evolution at the Crossroads: Modern Human Emergence in Western Asia

There is long-standing disagreement regarding Upper Pleistocene human evolution in Western Asia, particularly the Levant. Some argue that there were two dilierent populations, perhaps different species, of Upper Pleistocene Levantine hominids. The first, from the Israeli silcs of Qafzeh and Skhul. is anatomically modern. The second, from sites such as Amud. Kcbara. and Tabun, is archaic, or "Neandertal" in morphology. Others argue that ihis is a false dichotomy and that all of lliese hominids belong to a single, highly variable population. In this paper I attempt to resolve this issue by examining posteranial measures reflective of body shape. Results indicate that the Qafzeh-Skhul hominids have African-like, or tropically adapted, proportions, while tliosc from Amud, Kebara. Tabun. and Shanidar (Iraq) have more European-like, or cold-adapled. proportions. This suggests that iherc were in fact two distinct Western Asian populations and that the Qaf/ch-Skhul hominids were likely African in origin—i result consistent with the "Replacement"' model of modern human origins, [modern human origins, NeunJertals, Qafzeh-Skhul hominids, body shape]     

Dispersal and colonisation, long and short chronologies: how continuous is the Early Pleistocene record for hominids outside East Africa?

This paper examines the evidence for hominids outside East Africa during the Early Pleistocene. Most attention has focused recently on the evidence for or against a late Pliocene dispersal, ca. 1.8 Ma., of hominids out of Africa into Asia and possibly southern Europe. Here, the focus is widened to include North Africa as well as southern Asia and Europe, as well as the evidence in these regions for hominids after their first putative appearance ca. 1.8 Ma. It suggests that overall there is very little evidence for hominids in most of these regions before the Middle Pleistocene. Consequently, it concludes that the colonising capabilities of Homo erectus may have been seriously over-rated, and that even if hominids did occupy parts of North Africa, southern Europe and southern Asia shortly after 2 Ma, there is little evidence of colonisation. Whilst further fieldwork will doubtless slowly fill many gaps in a poorly documented Lower Pleistocene hominid record, it appears premature to conclude that the appearance of hominids in North Africa, Europe and Asia was automatically followed by permanent settlement. Rather, current data are more consistent with the view that Lower Pleistocene hominid populations outside East Africa were often spatially and temporally discontinuous, that hominid expansion was strongly constrained by latitude, and that occupation of temperate latitudes north of latitude 40 degrees was largely confined to interglacial periods.     

Inferring hominoid and early hominid phylogeny using craniodental characters: the role of fossil taxa

Recent discoveries of new fossil hominid species have been accompanied by several phylogenetic hypotheses. All of these hypotheses are based on a consideration of hominid craniodental morphology. However, Collard and Wood (2000) suggested that cladograms derived from craniodental data are inconsistent with the prevailing hypothesis of ape phylogeny based on molecular data. The implication of their study is that craniodental characters are unreliable indicators of phylogeny in hominoids and fossil hominids but, notably, their analysis did not include extinct species. We report here on a cladistic analysis designed to test whether the inclusion of fossil taxa affects the ability of morphological characters to recover the molecular ape phylogeny. In the process of doing so, the study tests both Collard and Wood's (2000) hypothesis of character reliability, and the several recently proposed hypotheses of early hominid phylogeny. One hundred and ninety-eight craniodental characters were examined, including 109 traits that traditionally have been of interest in prior studies of hominoid and early hominid phylogeny, and 89 craniometric traits that represent size-corrected linear dimensions measured between standard cranial landmarks. The characters were partitioned into two data sets. One set contained all of the characters, and the other omitted the craniometric characters. Six parsimony analyses were performed; each data set was analyzed three times, once using an ingroup that consisted only of extant hominoids, a second time using an ingroup of extant hominoids and extinct early hominids, and a third time excluding Kenyanthropus platyops. Results suggest that the inclusion of fossil taxa can play a significant role in phylogenetic analysis. Analyses that examined only extant taxa produced most parsimonious cladograms that were inconsistent with the ape molecular tree. In contrast, analyses that included fossil hominids were consistent with that tree. This consistency refutes the basis for the hypothesis that craniodental characters are unreliable for reconstructing phylogenetic relationships. Regarding early hominids, the relationships of Sahelanthropus tchadensis and Ardipithecus ramidus were relatively unstable. However, there is tentative support for the hypotheses that S. tchadensis is the sister taxon of all other hominids. There is support for the hypothesis that A. anamensis is the sister taxon of all hominids except S. tchadensis and Ar. ramidus. There is no compelling support for the hypothesis that Kenyanthropus platyops shares especially close affinities with Homo rudolfensis. Rather, K. platyops is nested within the Homo + Paranthropus + Australopithecus africanus clade. If K. platyops is a valid species, these relationships suggest that Homo and Paranthropus are likely to have diverged from other hominids much earlier than previously supposed. There is no support for the hypothesis that A. garhi is either the sister taxon or direct ancestor of the genus Homo. Phylogenetic relationships indicate that Australopithecus is paraphyletic. Thus, A. anamensis and A. garhi should be allocated to new genera.     

Faunal differences between the invasive brown macroalga Sargassum muticum and competing native macroalgae

Interactions between macroalgae and their associated fauna are of great interest for marine invasions, because fauna may increase the biotic resistance of a system and macroalgal invasions may cause shifts in faunal composition. We tested for differences in faunal community structure between a macroalgal invader, Sargassum muticum, and several native macroalgae in intertidal pools on both the west and south coast of Portugal. On each coast, we compared the faunal diversity and composition associated with the invader with that of the competing native macroalga(e). On the west coast, the diversity of the fauna associated with S. muticum was equal to or lower than with the native competitor, Cystoseira humilis. Fauna composition differed between S. muticum and C. humilis at both locations, but within each species, no differences between locations were detected. In contrast, the fauna diversity on S. muticum of the south coast varied among locations. S. muticum fauna differed from the fauna of all native macroalgae at one location, but only from three out of seven native macroalgae at the other location. Discriminating fauna species did not show a consistent pattern towards higher or lower abundances in S. muticum compared to most native macroalgae, and species-specific contributions were small. Differences in fauna community also depended on the identity of the native macroalga. In conclusion, the fauna associated with S. muticum differs from many native brown macroalgae, but these differences were not consistent as they depend both on the native macroalgal species and on location. This invader does not seem to have a severe negative impact on local macroalgae-associated fauna.     

Beyond leopards: tooth marks and the contribution of multiple carnivore taxa to the accumulation of the Swartkrans Member 3 fossil assemblage

The ca. 1.0 myr old fauna from Swartkrans Member 3 (South Africa) preserves abundant indication of carnivore activity in the form of tooth marks (including pits) on many bone surfaces. This direct paleontological evidence is used to test a recent suggestion that leopards, regardless of prey body size, may have been almost solely responsible for the accumulation of the majority of bones in multiple deposits (including Swartkrans Member 3) from various Sterkfontein Valley cave sites. Our results falsify that hypothesis and corroborate an earlier hypothesis that, while the carcasses of smaller animals may have been deposited in Swartkrans by leopards, other kinds of carnivores (and hominids) were mostly responsible for the deposition of large animal remains. These results demonstrate the importance of choosing appropriate classes of actualistic data for constructing taphonomic inferences of assemblage formation. In addition, they stress that an all-encompassing model of assemblage formation for the hominid-bearing deposits of the Sterkfontein Valley is inadequate and that each must be evaluated individually using not just analogical reasoning but also incorporating empirical data generated in the preserved fossil samples.     

The biological and chronological maturation of early hominids

Recent studies on the rate and pattern of dental development indicate that the growth and maturation of early hominids were more similar to the extant apes than to modern humans. This contrasts with the previously held opinion derived from combined dental development, pattern and attrition studies claiming that early hominids were more hominine in their development (Mann, 1975). This paper explores the origin of this difference of opinion and reviews immature hominid dentitions with the benefit of improved radiographs and new data on the pattern and rate of pongid dental development. Paranthropus and Australopithecus specimens are shown to possess an ape-like development pattern but incisor development is specialized in the former and superficially human-like in pattern. The present and recent studies on dental development rate and pattern justify the position that early hominids were more ape-like in their growth and development. Therefore, ages at death calculated from pongid dental development schedules are provided for most immature early hominids. More detailed studies of early hominid developmental biology are now possible. It is suggested that divergent heterochronic processes characterize changes in brain/body proportions during hominid evolution. Relative rates of bone remodeling processes can now be identified on early hominid skeletons. The paleodemographic analysis of early hominids is little changed by the developmental model one chooses.     

Meat-eating by early hominids at the FLK 22Zinjanthropussite, Olduvai Gorge (Tanzania): an experimental approach using cut-mark data

The meat-eating behavior of Plio-Pleistocene hominids, responsible for the bone accumulations at the earliest archaeological sites, is still a hotly-debated issue in paleoanthropology. In particular, meat-eating and bone marrow consumption are often presented as either complementary or opposing strategies of carcass exploitation. The presence of cut marks on fossil archeofauna is a potential source of information that has not been consistently used as evidence of carcass consumption by hominids. Some authors interpret cut marks as the result of hominids manipulating meat-bearing bones, while others argue that they can also be the result of hominids extracting marginal scraps of carcass flesh that have survived carnivores’ initial consumption. In this study, a referential framework concerning the interpretation of cut marks is presented, based on a set of experiments conducted by the author. It is suggested, according to these experiments and data drawn from the FLK “Zinj” site, that hominids processed meat-bearing bones (on which flesh was abundant) rather than defleshed carcasses from felid kills.     

北京凤凰山化石地点发掘简报

北京凤凰山地点位于周口店北京猿人遗址以北8km处, 是一处含哺乳动物化石与灰烬的洞穴-裂隙堆积。该地点包括两个化石层,上层为洞外的冲积物, 由棕红色黏土组成, 靠下部发现有零星的、破碎的化石, 可鉴定出似鸡骨山狐(Vulpes cf.chikushanensis)和羚羊(Antelopinae)两种类型; 下层含有灰烬层, 出土了较为丰富的哺乳动物化石, 可鉴定的哺乳动物化石包括鬣狗(Hyaenidae)、三门马(Equus sameniensis)、犀(Rhinocerotidae)、李氏野猪(Sus lydekkeri)、肿骨大角鹿(Megaloceros pachyosteus)、葛氏斑鹿(Cervus(Sika)grayi)、水牛(Bubalus sp.), 共7种。动物群与周口店第一地点和南京人化石地点的动物群相似, 指示该地点化石层的时代可能为中更新世。部分骨骼表面痕迹与灰烬层的发现显示该地点可能不排除有古人类活动痕迹的存在。     

Out of Africa: origins of the Taenia tapeworms in humans

Phylogenetic and divergence date analyses indicate that the occurrence of Taenia tapeworms in humans pre-dates the development of agriculture, animal husbandry and domestication of cattle (Bos spp.) or swine (Sus scrofa). Taeniid tapeworms in Africa twice independently colonized hominids and the genus Homo prior to the origin of modern humans. Dietary and behavioural shifts, from herbivory to scavenging and carnivory, as early Homo entered the carnivore guild in the Pliocene/Pleistocene, were drivers for host switching by tapeworms to hominids from carnivores including hyaenids and felids. Parasitological data provide a unique means of elucidating the historical ecology, foraging behaviour and food habits of hominids during the diversification of Homo spp.     

Human remains from Valdegoba Cave (Huérmeces, Burgos, Spain).

Systematic excavations, begun in 1987, at the Valdegoba cave site in northern Spain have yielded the remains of five individuals associated with a Middle Paleolithic stone tool technology and Pleistocene fauna. A fragmentary mandible of an adolescent (VB1), preserving nearly a full set of teeth, exhibits a symphyseal tubercle and slight incurvatio mandibulae anterior on the external symphysis. Both the superior and inferior transverse tori are present on the internal aspect. A second individual (VB2) is represented by a set of ten deciduous teeth consistent with an age at death of 6-9 months. A proximal manual phalanx (VB3) displays a relatively broad head, a characteristic which is found in both Neandertals, as well as European Middle Pleistocene hominids. VB4 is a fourth metatarsal that lacks the distal epiphysis, indicating it comes from an adolescent individual, and has a relatively high robusticity index. Finally, VB5 is a fifth metatarsal of an adult. The VB1 mandible shows a combination of archaic characteristics as well as more specific Neandertal morphological traits. The VB2 deciduous teeth are very small, and both the metrics and morphology seem more consistent with a modern human classification. The postcranial elements are undiagnostic, U-Th dating has provided an age of >350 ka for the base of the sequence and a date of <73.2+/-5 ka for level 7, near the top. Faunal analysis and radiometric dates from other nearby Mousterian sites suggests that the Valdegoba site is correlative with oxygen isotope stages 3-6 on the Iberian peninsula, and an Upper Pleistocene age for the Valdegoba hominids seems most reasonable.     

New estimates of tooth mark and percussion mark frequencies at the FLK Zinj site: the carnivore-hominid-carnivore hypothesis falsified

Traditional interpretations of hominid carcass acquisition strategies revolve around the debate over whether early hominids hunted or scavenged. A popular version of the scavenging scenario is the carnivore-hominid-carnivore hypothesis, which argues that hominids acquired animal resources primarily through passive opportunistic scavenging from felid-defleshed carcasses. Its main empirical support comes from the analysis of tooth mark frequency and distribution at the FLK Zinj site reported by Blumenschine (Blumenschine, 1995, J. Hum. Evol. 29, 21-51), in which it was shown that long bone mid-shafts exhibited a high frequency of tooth marks, only explainable if felids had preceded hominids in carcass defleshing. The present work shows that previous estimates of tooth marks on the FLK Zinj assemblage were artificially high, since natural biochemical marks were mistaken for tooth marks. Revised estimates are similar to those obtained in experiments in which hyenas intervene after humans in bone modification. Furthermore, analyses of percussion marks, notches, and breakage patterns provide data which are best interpreted as the results of hominid activity (hammerstone percussion and marrow extraction), based on experimentally-derived referential frameworks. These multiple lines of evidence support previous analyses of cut marks and their anatomical distribution; all indicate that hominids had early access to fleshed carcasses that were transported, processed, and accumulated at the FLK Zinj site.     

The evolution of brain size and intelligence in man

The brain size of hominids has increased approximately threefold during the evolution of the hominids fromAustralopithecus toHomo sapiens. It is proposed that the principal reason for this increase is that larger brains conferred greater intelligence, and greater intelligence conferred a selection advantage. A number of anthropologists have difficulty accepting this thesis because they believe that brain size is not associated with intelligence in man. Evidence is reviewed, and new evidence from two studies is presented, to show that brain size as measured by head size is positively correlated with intelligence as measured by intelligence tests. On two recent samples statistically significant correlations of .21 and .30 were obtained between estimates of brain size and IQ. It is considered that brain size is positively associated with intelligence in man and that this is the major reason for the increase in brain size of the hominids during the last 3.2 million years.