From apes to humans: locomotion as a key feature for phylogeny

If bipedalism has often been considered to be of a crucial interest for understanding hominid evolution, the acceptance of locomotor features to build phylogenies is still far from being a reality in the field. Especially for hominid evolution, it still seems to be difficult to accept that traits, other than craniodental ones, can be useful for defining the major dichotomies. The recent discovery of Australopithecus anamensis suggests a challenging view of the major dichotomy between apes and humans. Whilst it is widely accepted that Ardipithecus ramidus is ancestral to Australopithecus anamensis, which in its turn is ancestral to Australopithecus afarensis and then to later hominids, the postcranial adaptations, which should be taken into account, suggest another branching pattern. Based on the fact that by 4.0 million years two different locomotor patterns can be identified in hominids, two lineages would appear to be present: the "Australopithecine" lineage (with Australopithecus afarensis or Ardipithecus ramidus if the latter is really a hominid sensu stricto) and the "Hominine" lineage (with Australopithecus anamensis = Praeanthropus africanus).     

Research on late Pliocene Oldowan sites at Kanjera South, Kenya

The late Pliocene is notable for the appearance of two new hominid genera as well as the first archaeological sites, generally attributed to the Oldowan Industrial Complex. However, the behavioral ecology of Oldowan hominids has been little explored, particularly at sites older than 2.0 Ma. Moreover, debates on Oldowan hominid foraging ecology and behavior have centered on data from only two regions, and often from single site levels. Here we describe the preliminary results of our investigation of Oldowan occurrences at Kanjera South. These occurrences preserve the oldest known traces of hominid activity in southwestern Kenya, and unlike most of the Oldowan sites in the 2.0-2.5 Ma time interval, artefacts are found in spatial association with a well-preserved fauna. In 1996 and 1997, this project initiated the first excavation program for Kanjera South. Magneto- and biostratigraphy indicate that deposition began approximately 2.2 Ma, substantially earlier than previously thought. At Excavation 1, artefacts were found in spatial association with a taxonomically diverse faunal assemblage in Beds KS-1 and KS-2. Excavation 2 yielded a partial hippopotamus axial skeleton with artefacts in KS-3. Cores from both sites were incidentally flaked and represent a Mode I lithic technology indistinguishable from the Oldowan. Approximately 15% of the artefacts were manufactured from non-local raw materials, indicating a flow of resources into the area. Stable isotopic analysis of KS-1 and KS-2 pedogenic carbonates suggests that the Excavation 1 assemblages formed in a relatively open (>75% C4 grass) habitat. The Excavation 1 and 2 faunas contain a high proportion of equids relative to Oldowan accumulations from Bed I Olduvai Gorge, Tanzania. Beds KS-1 and KS-2 thus preserve traces of Oldowan hominid activities in a more open setting than has been previously documented.     

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.     

Reassessment of the paleoenvironment and preservation of hominid fossils from Hadar, Ethiopia.

Samples of paleosols from locality AL-333, known for numerous specimens of Australopithecus afarensis, were analyzed in order to reconstruct the original soils and environment of burial of the associated fossil hominids. The bones were found in swale-like features, within the calcareous and coarse-grained basal portion of a paleosol. This is more like an assemblage of bones buried during a single depositional episode, such as a flood, than an assemblage accumulated on a soil over a long period of time by carnivores or other means of death. What killed the hominids remains unclear, but considering the association of originally disarticulated bones of such hydraulically distinct types as phalanges and maxillae, it is very likely that they died and partially rotted at or very near this site. The paleosols at AL-333, here named the Fo and Go clay paleosols, have calcareous rhizoconcretions, granular surface horizons, prismatic peds, and shallow calcareous nodules and stringers like soils now supporting grassy woodland in semiarid regions. Although this group of hominids was buried in streamside gallery woodland, there is evidence from Laetoli, Tanzania, that A. afarensis ventured out into open wooded grassland as well. Evidence for this should be sought from other paleosols at Hadar.     

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.     

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.     

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.     

Phylogeny of early Australopithecus: new fossil evidence from the Woranso-Mille (central Afar,Ethiopia)

The earliest evidence of Australopithecus goes back to ca 4.2 Ma with the first recorded appearance of Australopithecus ‘anamensis’ at Kanapoi, Kenya. Australopithecus afarensis is well documented between 3.6 and 3.0 Ma mainly from deposits at Laetoli (Tanzania) and Hadar (Ethiopia). The phylogenetic relationship of these two ‘species’ is hypothesized as ancestor–descendant. However, the lack of fossil evidence from the time between 3.6 and 3.9 Ma has been one of its weakest points. Recent fieldwork in the Woranso-Mille study area in the Afar region of Ethiopia has yielded fossil hominids dated between 3.6 and 3.8 Ma. These new fossils play a significant role in testing the proposed relationship between Au. anamensis and Au. afarensis. The Woranso-Mille hominids (3.6–3.8 Ma) show a mosaic of primitive, predominantly Au. anamensis-like, and some derived (Au. afarensis-like) dentognathic features. Furthermore, they show that, as currently known, there are no discrete and functionally significant anatomical differences between Au. anamensis and Au. afarensis. Based on the currently available evidence, it appears that there is no compelling evidence to falsify the hypothesis of ‘chronospecies pair’ or ancestor–descendant relationship between Au. anamensis and Au. afarensis. Most importantly, however, the temporally and morphologically intermediate Woranso-Mille hominids indicate that the species names Au. afarensis and Au. anamensis do not refer to two real species, but rather to earlier and later representatives of a single phyletically evolving lineage. However, if retaining these two names is necessary for communication purposes, the Woranso-Mille hominids are best referred to as Au. anamensis based on new dentognathic evidence.     

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.     

Species-specific albumin in fossil bones from Orce, Granada, Spain

A skull fragment (VM-0) from Orce, Granada, Spain, dated palaeomagnetically at about 1.6 Myr, is thought by some palaeontologist to be hominid, while others maintain it is equid. If hominid, it would be by far the oldest evidence ofHomo in Europe. Immunological studies on residual albumin in this fossil were carried out independently, and with different immunological methods, at the University of California, San Francisco (radioimmunoassay), and at the University of Granada, Spain (enzyme immunoassay). Other fossils attributed to hominids also studied wereVM1960 from Venta Micena, andCV-1 from Cueva Victoria, Murcia, Spain. Undisputed equid and bovid fossils from the same deposits and dated to a similar period as the Orce skull were also analyzed. Our results showed that species-specific albumin can be detected in 1.6 Myr-old hominid, equid and bovid fossils. The albumin from the Orce skull fragment and fromVM-1960 was immunologically closer to human albumin. These findings support the contention that theVM-0 andVM-1960 are hominid and that members of the genusHomo occupied southern Spain 1.6 Myr ago.     

Physiology, thermoregulation and bipedalism

It has long been recognized that the bipedal posture reduces the surface area of the body exposed to the sun. In recent years, a theory has been developed by Wheeler that bipedalism evolved in the ancestor of the Hominidae in order to help relieve thermal stress on the animals in open equatorial environments. Bipedalism was said to afford a distinct adaptive advantage over quadrupedalism by permitting hominids to remain active in the open throughout the day. The heat load of the hypothetical hominid comprises the external environment as modelled by Wheeler and the animal's internal environment (i.e., the internal heat generated by its metabolic and locomotor activities, and its evaporative and respirative cooling capacities). When these factors are integrated in the calculation of the animal's thermal budget, the putative advantage of the bipedal over the quadrupedal posture is considerably reduced. The simulations conducted in this study suggest that the increased time afforded to early hominids in the open by bipedalism was relatively short and, therefore, of little or no adaptive significance. These results suggest that thermoregulatory considerations cannot be implicated as a first cause in the evolution of bipedalism in the hominid ancestor.     

The geological context of the Kanapoi fossil hominids

Recent study of the geological succession at Kanapoi reveals that there are at least three series of sediments younger than the early Pliocene Kanapoi sediments which repose unconformably on them. Both sets of terrace and placage deposits contain an admixture of reworked Pliocene fossils and younger fossils preserved at the time of deposition of the younger sediments. This discovery throws doubt on the homogeneous nature of the Kanapoi fossil hominid sample, and suggests instead thatAustralopithecus anamensis may consist of a chimera of an early Pliocene hominid with generally ape-like dentognathic and postcranial anatomy and considerably youngerHomo specimens with more human-like post-cranial bones.     

Influence of Plio-Pleistocene aridification on human evolution: evidence from paleosols of the Turkana Basin, Kenya

New stable carbon isotope measurements, coupled with paleoprecipitation estimates, both from Plio-Pleistocene paleosols of the Turkana Basin, Kenya, provide a high-resolution record of aridification and increasing C4 biomass during the past 4.3 Ma. This aridification trend is marked by several punctuations at 3.58-3.35, 2.52-2, and 1.81-1.58 Ma, during which the running mean and variance of delta13C and paleoaridity estimates increase, suggesting that the proportion of C4 biomass increases in savanna mosaics during periods of heightened aridity. Increase in C4 biomass during these aridification events not only increases the proportion of open habitats, but increases the spatial neg-entropy, or heterogeneity of the ecosystem. The aridification events identified correspond to intervals of increased turnover, but more importantly, increased diversity of bovids. Although the record of hominins from the Turkana Basin lacks the temporal resolution and diversity of the bovid record, the aridification intervals identified are marked by similar increases in the diversity and turnover of hominins. These results support the hypothesis that hominins evolved in savanna mosaics that changed through time, and suggest that the evolution of bovids and hominins was driven by shifts in climatic instability and habitat variability, both diachronic and synchronic.     

Endocranial hyperostosis in Sangiran 2, Gibraltar 1, and Shanidar 5

Sangiran hominid 2 (S-2), Gibraltar hominid 1 (G-1), and Shanidar hominid 5 (SH-5) exhibit previously undescribed bilateral, paramedian hyperostosis of the endocranial frontal squama that spares the frontal crest, sagittal sinus, and ectocranial surface. The hyperostosis is localized to the frontal (usually the middle third) and parietal and is consistent with a diagnosis of hyperostosis calvariae interna (HCI), inclusive of hyperostosis frontalis interna. The hyperostosis in these specimens is compared to fossil hominids from Indonesia and Europe and to modern human cases of HCI. The three cases of HCI reported here document the existence and frequency of HCI in fossil hominids and push the antiquity of the disease back to nearly 1.5 millin years. The relatively great incidence of HCI in fossil hominids adds another confounding factor to the problematical issue of the taxonomic significance of cranial vault thickness. Am J Phys Anthropol 102:111–122. © 1997 Wiley-Liss, Inc.     

Was Australopithecus anamensis ancestral to A. afarensis? A case of anagenesis in the hominin fossil record

We tested the hypothesis that early Pliocene Australopithecus anamensis was ancestral to A. afarensis by conducting a phylogenetic analysis of four temporally successive fossil samples assigned to these species (from earliest to latest: Kanapoi, Allia Bay, Laetoli, Hadar) using polarized character-state data from 20 morphological characters of the dentition and jaws. If the hypothesis that A. anamensis is ancestral to A. afarensis is true, then character-state changes between the temporally ordered site-samples should be congruent with hypothesized polarity transformations based on outgroup (African great ape) conditions. The most parsimonious reconstruction of character-state evolution suggests that each of the hominin OTUs shares apomorphies only with geologically younger OTUs, as predicted by the hypothesis of ancestry (tree length=31; Consistency Index=0.903). This concordance of stratigraphic and character-state data supports the idea that the A. anamensis and A. afarensis samples represent parts of an anagenetically evolving lineage, or evolutionary species. Each site-sample appears to capture a different point along this evolutionary trajectory. We discuss the implications of this conclusion for the taxonomy and adaptive evolution of these early-middle Pliocene hominins.     

Biomechanical interpretation of the early hominid hip

A new pelvic fragment from Swartkrans provides the opportunity to analyze the hip joint mechanics of the robust form of early hominid. The function of the lateral support system provided by the abductor muscles of the hip appears to be similar to that of the gracile early hominid from Sterkfontein. The system is well adapted for providing the lateral support necessary for efficient bipedalism. The hip extensor mechanism and hip internal rotatory system also appear to be well adapted for efficient bipedalism in a way very similar to the other early hominids. The conclusion reached is that the robust and gracile forms of South African early hominids were basically similar in their locomotor adaptation and were most likely habitual bipeds.     

Neanderthal Skeletal Structure and the Place of Homo neanderthalensis in European Hominid Phylogeny

Although the debate rages on over whether the Neanderthals merit their own species status or should be viewed as an odd variant of Homo sapiens, recent evidence has accumulated that overwhelmingly supports the former interpretation. Among this evidence is a recent full-body skeletal reconstruction that not only highlights the extreme differences between the highly apomorphic H. sapiens and H. neanderthalensis in the construction of the thorax and pelvic girdle, but strongly suggests significant gait differences between the two species that add to the probability that the two kinds of hominid would not have recognized each other as breeding partners. This is hardly surprising since the two species possessed a relatively remote common ancestry, and it is indeed suggested here that Homo neanderthalensis was merely one species embedded within a diverse and endemic middle Pleistocene European hominid radiation. Clearly more than one lineage of hominids simultaneously occupied Europe during the middle Pleistocene.     

Paleoenvironmental reconstruction of middle Miocene paleosols bearing Kenyapithecus and Victoriapithecus, Nyakach Formation, southwestern Kenya

Paleosols in the middle Miocene (15 Ma) Nyakach Formation at Kaimogool, near Sondu, southwestern Kenya have yielded specimens of the early cercopithecoid Victoriapithecus macinnesi and the early kenyapithecine Kenyapithecus africanus, and can be used as evidence for the environmental mosaic occupied by these primates. Five distinct types of paleosols (pedotypes) are recognized in the Nyakach Formation section at Kaimogool South. The most common paleosols are reddish brown, silty calcareous profiles with blocky structure and large root traces (Ratong pedotype) which are interpreted as soils of well-drained, dry bushland or thicket (nyika). Weakly developed paleosols associated with paleochannels (Dhero pedotype) represent wooded grassland early in the ecological succession from streamside flooding. One of these paleosols has yielded a fossil flora of grasses and small-leaved dicots like those of modern semi-arid wooded grassland. Crumb structured, calcareous paleosols with iron-manganese nodules (Yom pedotype) are interpreted to represent seasonally waterlogged, wooded grassland (dambo or vlei). Thick, red clayey, calcareous paleosols with blocky ped structure and large root traces (Tut pedotype) are interpreted as soils of well-drained dry woodland. Other blocky-structured, gray to brown calcareous paleosols with iron-manganese nodules (Chido pedotype) are interpreted as soils of seasonally waterlogged, riparian dry woodland. Fossil soils, plants and gastropods are evidence of an unusually dry (300-500 mm mean annual precipitation) habitat for apes, consisting of a vegetational mosaic dominated by dry woodland, bushland and thickets with few areas of seasonally waterlogged grassland. Fossils of V. macinnesi are rare from Nyakach, but were found in paleosols representative of bushland and thicket habitats (Ratong). Fossils of the ape K. africanus were found within paleosols indicative of dry woodland (Tut). Other paleosol types representative of seasonally dry dambo grassland (Yom), colonizing grassland (Dhero) or riparian woodland (Chido) are also represented, but have not yet produced primate fossils.     

化石人类脑演化研究概况

脑演化是人类演化的一个重要组成部分,其研究可以为人类起源、演化、人群关系及语言、智力等方面提供重要的信息。脑演化的主要证据是通过研究颅内模（endocast）及颅骨的形态得到的。颅内模是从颅骨内表面得到的脑的外部形态。有时颅骨的内腔充满泥沙,并且为钙质所结固,可以自然形成颅内模。也可以人工制作颅内模。颅内模和颅骨的内表面能够提供人类脑髓及神经进化方面的直接证据。对化石人类脑演化的研究主要包括以下几个方面的内容：测量或估计脑量的大小及其和身体大小之间的关系;研究脑量随时间的变化过程;通过对早期人类颅内模表面沟回形态特征的研究,探索脑功能区在早期人类和猿类的区别及在演化上的变化;左右大脑不对称性与一侧优势关系,探讨语言的起源和惯用手的脑功能基础等;脑膜中动脉系统、静脉窦系统及与血液循环相关的排泄孔的变化,探讨大脑各部分比例的变化和功能的日益复杂对供血需求的影响;通过对人类脑演化的研究,探讨人类进化的原因。本文通过对以上几个方面及其中国化石脑演化研究的介绍,对化石人类脑演化的研究概况作综合论述和简单回顾。