The Middle Stone Age archaeology of the Lower Omo Valley Kibish Formation: excavations, lithic assemblages, and inferred patterns of early Homo sapiens behavior

This paper describes the excavation, stratigraphy, and lithic assemblages of Middle Stone Age sites from the Omo Kibish Formation (Lower Omo Valley, southwestern Ethiopia). Three sites were excavated, two in Kibish Member I (KHS and AHS) and one at the base of Member III (BNS). The assemblages are dominated by relatively high-quality raw materials procured as pebbles from local gravels. The principal modes of core preparation are radial/centripetal Levallois and discoidal. Retouched tools are rare. Foliate bifaces are present, as are larger tools, such as handaxes, picks, and lanceolates, but these are more common among surface finds than among excavated assemblages. Middle Stone Age assemblages shed light on the adaptations of the earliest-known Homo sapiens populations in Africa.     

Microstratigraphy of the Kibish hominin sites KHS and PHS, Lower Omo Valley, Ethiopia

Detailed stratigraphic analysis of the Omo I and Omo II fossil localities confirms both the relational and sequential context reported by Butzer in 1969. The two fossils derive from approximately the same level within upper Member I of the Kibish Formation. Additional features of the local stratigraphic sequences indicate a complex history of depositional events, minor erosional surfaces, and weak soil formation throughout upper Member I.     

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.     

A description of the Omo I postcranial skeleton, including newly discovered fossils

Recent fieldwork in the Kibish Formation has expanded our knowledge of the geological, archaeological, and faunal context of the Omo I skeleton, the earliest known anatomically modern human. In the course of this fieldwork, several additional fragments of the skeleton were recovered: a middle manual phalanx, a distal manual phalanx, a right talus, a large and a small fragment of the left os coxae, a portion of the distal diaphysis of the right femur that conjoins with the distal epiphysis recovered in 1967, and a costal fragment. Some researchers have described the original postcranial fragments of Omo I as anatomically modern but have noted that a variety of aspects of the specimen's morphology depart from the usual anatomy of many recent populations. Reanalysis confirms this conclusion. Some of the unusual features in Omo I--a medially facing radial tuberosity, a laterally flaring facet on the talus for the lateral malleolus, and reduced dorsovolar curvature of the base of metacarpal I--are shared with Neandertals, some early modern humans from Skhul and Qafzeh, and some individuals from the European Gravettian, raising the possibility that Eurasian early modern humans inherited these features from an African predecessor rather than Neandertals. The fragment of the os coxae does not unambiguously diagnose Omo I's sex: the greater sciatic notch is intermediate in form, the acetabulum is large (male?), and a preauricular sulcus is present (female?). The preserved portion of the left humerus suggests that Omo I was quite tall, perhaps 178-182 cm, but the first metatarsal suggests a shorter stature of 162-173 cm. The morphology of the auricular surface of the os coxae suggests a young adult age.     

Sapropels and the age of hominins Omo I and II, Kibish, Ethiopia

The provenance and age of two Homo sapiens fossils (Omo I and Omo II) from the Kibish Formation in southern Ethiopia have been much debated. Here we confirm that Omo I and the somewhat more primitive-looking Omo II calvariae are from similar stratigraphic levels in Member I of the Kibish Formation. Based on (40)Ar/(39)Ar age measurements on alkali feldspar crystals from pumice clasts in the Nakaa'kire Tuff, a tuffaceous bed in Member I just below the hominin levels, we place an older limit of 198+/-14ka (weighted mean age=196+/-2ka) for the hominins. A younger limit of 104+/-7ka (weighted mean age=104+/-1ka) is provided by feldspars separated from pumice clasts in the Aliyo Tuff in Member III. Geological evidence indicates rapid deposition of each member of the Kibish Formation, concurrent with deposition of sapropels in the Mediterranean Sea. The (40)Ar/(39)Ar age measurements, together with correlations with sapropels, indicate that the hominin fossils are close in age to the older limit. Our preferred estimate of the age of the hominins is 195+/-5ka, making them the earliest well-dated anatomically modern humans yet described.     

Stratigraphy and tephra of the Kibish Formation, southwestern Ethiopia

The Kibish Formation in southwestern Ethiopia, with an aggregate thickness of ~105m, consists of lacustrine, marginal lacustrine, and deltaic deposits. It is divided into four members numbered I to IV on the basis of erosion surfaces (disconformities) between the strata of each member. It overlies the Mursi and Nkalabong formations, the latter of which is here shown to correlate with the Shungura Formation. Tephra layers in each member allow for secure correlation between geographically separated sections on the basis of the composition of their volcanic glass. Members I, III, and IV of the Kibish Formation appear to have been deposited at the same times as sapropels S7 (197ka), S4 (104ka), and S1 (8ka) in the eastern Mediterranean Sea, respectively. We correlate the KHS Tuff of the Kibish Formation with a >154-kyr-old unnamed tuff in the Konso Formation. Tephra in Member IV may derive from Mount Wenchi, a volcano situated on the divide between the Omo and Blue Nile drainage basins. Thin-bedded sedimentary layers probably represent annual deposition reflecting rapid sedimentation (~30m/kyr) of parts of the formation. This conclusion is supported by variation in paleomagnetic inclination through a sequence of these layers at KHS. Two fossils of early Homo sapiens (Omo I and Omo II) derive from Member I. Their stratigraphic placement is confirmed by analysis of the KHS Tuff in the lower part of Member II at both fossil sites. The KHS Tuff lies above a disconformity, which itself lies above the fossils at both sites. (40)Ar/(39)Ar dates provide an estimated age of ~195kyr for these fossils. Omo III, a third fossil H. sapiens, probably also derives from Member I of the Kibish Formation and is of similar age. Hominin fossils from AHS, a new site, also derive from Member I. Hominin fossils from CHS can only be placed between 104ka and 10ka, the H. sapiens specimen from JHS is most likely 9-13kyr in age, and a partial skeleton of H. sapiens from Pelvic Corner is most likely ~6.6kyr in age.     

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.     

Dental microwear and diets of African early Homo

Conventional wisdom ties the origin and early evolution of the genus Homo to environmental changes that occurred near the end of the Pliocene. The basic idea is that changing habitats led to new diets emphasizing savanna resources, such as herd mammals or underground storage organs. Fossil teeth provide the most direct evidence available for evaluating this theory. In this paper, we present a comprehensive study of dental microwear in Plio-Pleistocene Homo from Africa. We examined all available cheek teeth from Ethiopia, Kenya, Tanzania, Malawi, and South Africa and found 18 that preserved antemortem microwear. Microwear features were measured and compared for these specimens and a baseline series of five extant primate species (Cebus apella, Gorilla gorilla, Lophocebus albigena, Pan troglodytes, and Papio ursinus) and two protohistoric human foraging groups (Aleut and Arikara) with documented differences in diet and subsistence strategies. Results confirmed that dental microwear reflects diet, such that hard-object specialists tend to have more large microwear pits, whereas tough food eaters usually have more striations and smaller microwear features. Early Homo specimens clustered with baseline groups that do not prefer fracture resistant foods. Still, Homo erectus and individuals from Swartkrans Member 1 had more small pits than Homo habilis and specimens from Sterkfontein Member 5C. These results suggest that none of the early Homo groups specialized on very hard or tough foods, but that H. erectus and Swartkrans Member 1 individuals ate, at least occasionally, more brittle or tough items than other fossil hominins studied.     

Estimation of the Extent of Synteny Between Tetraodon nigroviridis and Homo sapiens Genomes

This paper presents a genomic comparison between 20 sequenced BACs (or fragments of BACs) from Tetraodon nigroviridis and the human genome. A total of 199 fish genes were identified by informatics resources, together with their putative human orthologues. Comparisons of the localizations in both species led to the identification of 32 syntenic regions and a minimum of 131 rearrangements in these regions that occurred during independent evolution of these species. This made it possible to estimate the rate of genomic rearrangements that occurred per million years (and per megabase). This rate is comparable to that obtained by comparison of the Fugu rubripes shotgun sequence data to human data but is significantly higher that those obtained by comparing the human genome to mammalian genomes. Overall, it suggests that genomic evolution by rearrangement is not uniform within the vertebrate group.Sequence data for the genomic BAC clones have been deposited with the DDBJ/EMBL/GenBank Data Libraries under accession numbers BX629360, BX629354, BX629355, BX629356, BX629357, BX629358, BX629359, and BX629360.     

Energetics in Homo erectus and other early hominins: the consequences of increased lower-limb length

Previous studies of daily energy expenditure (DEE) in hominin fossils have estimated locomotor costs using a formula that was based on six species, all 18 kg or less in mass, including no primates, and that has a number of other problems when applied in an ecological context. It is well established that the energetic cost of human walking is lower than that of representative mammals, particularly for individuals with long lower limbs. The current study reevaluates the daily energy expenditures of a variety of hominin species using more appropriate approaches to estimating locomotor costs. To estimate DEE for primates, I relied on published data on body mass, day range, and the percentage of time spent in various activities. Based on those data, I calculated a value for nonlocomotor DEE. I then used a variant of a method that I have suggested elsewhere to calculate the daily cost due to locomotion (DEEL) and summed the two to calculate total DEE. The more up-to-date methods for calculating the cost of travel result in lower estimates of this aspect of the energy budget than seen in previous studies. Values obtained here for DEE in various representatives of Australopithecus are lower than reported previously by around 200 kcal/day. Taking into account the greater economy of human walking, particularly the effect of the longer lower limbs found in many later Homo species, also results in lowered estimates of DEE. Elongation of the lower limbs in H. erectus reduced relative travel costs nearly 50% in comparison to A.L. 288-1 (A. afarensis). The present method for calculating DEE indicates that female H. erectus DEE was 84% greater than that of female Australopithecus; this disparity is even larger than that suggested by previous workers.     

Cladistic analysis of early Homo crania from Swartkrans and Sterkfontein, South Africa

The phylogenetic relationships of early Pleistocene Homo crania from the South African sites of Swartkrans and Sterkfontein were investigated through cladistic analyses of 99 morphological characters. The Swartkrans Member 1 specimen SK 847 and the Stw 53 cranium from Sterkfontein Member 5A were treated as separate operational taxonomic units (OTUs), distinct from the three species of early Homo-H. erectus, H. habilis, and H. rudolfensis-that are recognized from the Plio-Pleistocene deposits of East Africa. The cladistic analyses differed in the treatment of the South African OTUs (separate Swartkrans and Sterkfontein OTUs vs. a single Swartkrans+Sterkfontein OTU). PAUP 4.0 was used to construct cladograms and address hypotheses about relationships. In the analysis that treated the South African specimens as a single OTU, the position of that OTU was stable as a separate branch on the Homo clade between H. rudolfensis and [H. habilis+(H. erectus+H. sapiens)]. When SK 847 and Stw 53 were treated as separate OTUs, the majority of most parsimonious trees indicated that they were positioned in similar positions as the combined South African Homo OTU; that is, as separate branches between H. rudolfensis and [H. habilis+(H. erectus+H. sapiens)], with the Swartkrans OTU generally occupying a more derived position. The position of the Sterkfontein OTU was more stable than that of the Swartkrans OTU, which was found in several other positions among the minimum length trees. Running the analyses with only those characters preserved by SK 847 and Stw 53 resulted in similar topologies for minimum length trees, although the positions of Stw 53, SK 847, and H. habilis exchanged places in some trees. In no case was an exclusive sister relationship between either South African OTU and a particular species of Homo supported statistically. Both South African OTUs differ from H. habilis in the fewest number of cladistic characters.     

Liang Bua Homo floresiensis mandibles and mandibular teeth: a contribution to the comparative morphology of a new hominin species

In 2004, a new hominin species, Homo floresiensis, was described from Late Pleistocene cave deposits at Liang Bua, Flores. H. floresiensis was remarkable for its small body-size, endocranial volume in the chimpanzee range, limb proportions and skeletal robusticity similar to Pliocene Australopithecus, and a skeletal morphology with a distinctive combination of symplesiomorphic, derived, and unique traits. Critics of H. floresiensis as a novel species have argued that the Pleistocene skeletons from Liang Bua either fall within the range of living Australomelanesians, exhibit the attributes of growth disorders found in modern humans, or a combination of both. Here we describe the morphology of the LB1, LB2, and LB6 mandibles and mandibular teeth from Liang Bua. Morphological and metrical comparisons of the mandibles demonstrate that they share a distinctive suite of traits that place them outside both the H. sapiens and H. erectus ranges of variation. While having the derived molar size of later Homo, the symphyseal, corpus, ramus, and premolar morphologies share similarities with both Australopithecus and early Homo. When the mandibles are considered with the existing evidence for cranial and postcranial anatomy, limb proportions, and the functional anatomy of the wrist and shoulder, they are in many respects closer to African early Homo or Australopithecus than to later Homo. Taken together, this evidence suggests that the ancestors of H. floresiensis left Africa before the evolution of H. erectus, as defined by the Dmanisi and East African evidence.     

Who made the Aurignacian and other early Upper Paleolithic industries?

The Aurignacian is typically taken as a marker of the spread of anatomically modern humans into Europe. However, human remains associated with this industry are frustratingly sparse and often limited to teeth. Some have suggested that Neandertals may, in fact, be responsible for the Aurignacian and the earliest Upper Paleolithic industries. Although dental remains are frequently considered to be taxonomically undiagnostic in this context, recent research shows that Neandertals possess a distinct dental pattern relative to anatomically modern humans. Even so, it is rare to find mandibles or maxillae that preserve all or most of their teeth; and, the probability of correctly identifying individuals represented by only a few teeth or a single tooth is unknown. We present a Bayesian statistical approach to classifying individuals represented exclusively by teeth into two possible groups. The classification is based on dental trait frequencies and sample sizes for ‘known’ samples of 95 Neandertals and 63 Upper Paleolithic modern humans. In a cross validation test of the known samples, 89% of the Neandertals and 89% of the Upper Paleolithic modern humans were classified correctly. We then classified an ‘unknown’ sample of 52 individuals: 34 associated with Aurignacian or other (non-Châtelperronian) early Upper Paleolithic industries, 15 associated with the Châtelperronian, and three unassociated. Of the 34 early Upper Paleolithic-associated individuals, 29 were assigned to modern humans, which is well within the range expected (95% of the time 26-33) with an 11% misclassification rate for an entirely modern human sample. These results provide some of the strongest evidence that anatomically modern humans made the Aurignacian and other (non-Châtelperronian) early Upper Paleolithic industries.     

Human talus bones from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca,Burgos, Spain)

Here we present and describe comparatively 25 talus bones from the Middle Pleistocene site of the Sima de los Huesos (SH) (Sierra de Atapuerca, Burgos, Spain). These tali belong to 14 individuals (11 adult and three immature). Although variation among Middle and Late Pleistocene tali tends to be subtle, this study has identified unique morphological characteristics of the SH tali. They are vertically shorter than those of Late Pleistocene Homo sapiens, and show a shorter head and a broader lateral malleolar facet than all of the samples. Moreover, a few shared characters with Neanderthals are consistent with the hypothesis that the SH population and Neanderthals are sister groups. These shared characters are a broad lateral malleolar facet, a trochlear height intermediate between modern humans and Late Pleistocene H. sapiens, and a short middle calcaneal facet. It has been possible to propose sex assignment for the SH tali based on their size. Stature estimates based on these fossils give a mean stature of 174.4 cm for males and 161.9 cm for females, similar to that obtained based on the long bones from this same site.     

People of the ancient rainforest: late Pleistocene foragers at the Batadomba-lena rockshelter, Sri Lanka

Batadomba-lena, a rockshelter in the rainforest of southwestern Sri Lanka, has yielded some of the earliest evidence of Homo sapiens in South Asia. H. sapiens foragers were present at Batadomba-lena from ca. 36,000 cal BP to the terminal Pleistocene and Holocene. Human occupation was sporadic before the global Last Glacial Maximum (LGM). Batadomba-lena’s Late Pleistocene inhabitants foraged for a broad spectrum of plant and mainly arboreal animal resources (monkeys, squirrels and abundant rainforest snails), derived from a landscape that retained equatorial rainforest cover through periods of pronounced regional aridity during the LGM. Juxtaposed hearths, palaeofloors with habitation debris, postholes, excavated pits, and animal and plant remains, including abundant Canarium nutshells, reflect intensive habitation of the rockshelter in times of monsoon intensification and biome reorganisation after ca. 16,000 cal BP. This period corresponds with further broadening of the economic spectrum, evidenced though increased contribution of squirrels, freshwater snails and Canarium nuts in the diet of the rockshelter occupants. Microliths are more abundant and morphologically diverse in the earliest, pre-LGM layer and decline markedly during intensified rockshelter use on the wane of the LGM. We propose that changing toolkits and subsistence base reflect changing foraging practices, from shorter-lived visits of highly mobile foraging bands in the period before the LGM, to intensified use of Batadomba-lena and intense foraging for diverse resources around the site during and, especially, following the LGM. Traces of ochre, marine shell beads and other objects from an 80 km-distant shore, and, possibly burials reflect symbolic practices from the outset of human presence at the rockshelter. Evidence for differentiated use of space (individual hearths, possible habitation structures) is present in LGM and terminal Pleistocene layers. The record of Batadomba-lena demonstrates that Late Pleistocene pathways to (aspects of) behavioural ‘modernity’ (composite tools, practice of symbolism and ritual, broad spectrum economy) were diverse and ecologically contingent.     

Palaeopathological and variant conditions of the Homo heidelbergensis type specimen (Mauer,Germany)

Although early Homo specimens are now known from a number of African, Asian and European Middle Pleistocene sites, the taxon Homo heidelbergensis was initially introduced for the Mauer jaw recovered in 1907. Fossil hominids from the earlier Middle Pleistocene of Europe are very rare and the Mauer mandible is generally accepted as one of the most ancient, with an age of approximately 700 kyr. A new preparation of the mandible was conducted in 1996 and gave rise to the detailed palaeopathological examination which is presented here. Based on comparative analyses, the extreme breadth of the mandibular ramus and its flat intercondylar incision, in conjunction with the flattening and broadening of the coronoid process tip, results either from an idiosyncratic pattern of the course and insertion of the temporalis muscle on the coronoid process or from the temporalis possessing an accessory head. The incidence of periodontal pocketing, together with a vertical reduction of the alveolar margin to approximately 3.00 mm, and a slight protuberance formed in vicinity of the right M(2)can safely be interpreted as pathognomonic indications of periodontal disease. The short distance between the enamel-dentine junction of the teeth and the horizontal alveolar margins could either be an inherited variant or may result from incipient osteoporosis. In addition, an arthrotic condition with slight osteophytic peripheral exostoses and an arthrolit (i.e. an articular calculus or "joint mouse") on the left condylus articularisand a depression in the medial part of the left mandibular condyle extending into the inferior part of the ramus are present. These features are indicative of a trauma-induced osteochondrosis dissecans. The diagnosis therefore suggests that the observed depression results from a well-healed fracture. This traumatic event illustrates the demanding living conditions endured by humans during the European Middle Pleistocene. The variations and pathological conditions observed in Mauer do not question the mandible's role as type specimen for the taxon Homo heidelbergensis.     

Structural and kinetic characterization of quinolinate phosphoribosyltransferase (hQPRTase) from homo sapiens

Human quinolinate phosphoribosyltransferase (EC 2.4.2.19) (hQPRTase) is a member of the type II phosphoribosyltransferase family involved in the catabolism of quinolinic acid (QA). It catalyses the formation of nicotinic acid mononucleotide from quinolinic acid, which involves a phosphoribosyl transfer reaction followed by decarboxylation. hQPRTase has been implicated in a number of neurological conditions and in order to study it further, we have carried out structural and kinetic studies on recombinant hQPRTase. The structure of the fully active enzyme overexpressed in Escherichia coli was solved using multiwavelength methods to a resolution of 2.0 A. hQPRTase has a alpha/beta barrel fold sharing a similar overall structure with the bacterial QPRTases. The active site of hQPRTase is located at an alpha/beta open sandwich structure that serves as a cup for the alpha/beta barrel of the adjacent subunit with a QA binding site consisting of three arginine residues (R102, R138 and R161) and two lysine residues (K139 and K171). Mutation of these residues affected substrate binding or abolished the enzymatic activity. The kinetics of the human enzyme are different to the bacterial enzymes studied, hQPRTase is inhibited competitively and non-competitively by one of its substrates, 5-phosphoribosylpyrophosphate (PRPP). The human enzyme adopts a hexameric arrangement, which places the active sites in close proximity to each other.