A revised stratigraphic framework for Olduvai Gorge Bed I based on tuff geochemistry

Olduvai Gorge, Tanzania has rich records of Plio-Pleistocene fauna and flora, hominin fossils, and stone artifacts preserved between well-dated tephra layers (tuffs). Accurate correlation between sites in the two million year section is complicated by faulting, erosion, change in physical appearance of the tuffs, and quality of preservation. Traditional tuff geochemical techniques using glass cannot be applied because of poor glass preservation, and previous physical mapping has led to miscorrelations in Bed I. A new approach, using a combination of glass and mineral compositions (feldspar, augite, hornblende, and oxides) produced successful geochemical fingerprints for all ten major Bed I (∼2.03-1.79 Ma) tuffs. These fingerprints make available a reliable means for correlating specific tuffs between the well-dated "Junction" sites, such as FLK and HWK, and less well-dated sites at the basin margins. The new correlations provide a high-resolution stratigraphic framework for Bed I and correct previous miscorrelations in the west of the basin. Olduvai Hominin 65, from western Olduvai, was recovered from a level between Tuff IC and the Ng’eju Tuff, and therefore dates to 1.79-1.84 Ma.     

A Homo habilis maxilla and other newly-discovered hominid fossils from Olduvai Gorge, Tanzania

In 1995, a 1.8 million year old hominid maxilla with complete dentition (OH 65) was excavated from Bed I in the western part of Olduvai Gorge. The molar crowns are small relative to the long flaring roots, and the root of the canine is very long and straight. The broad maxilla with wide U-shaped palate and the form of the tooth roots closely match those of KNM-ER 1470 which, in its parietal size and morphology, matches the type specimen of Homo habilis, OH 7. Thus, OH 65 and KNM-ER 1470 group with OH 7 as representatives of H. habilis while some other Olduvai specimens, such as OH 13 and OH 24, have more in common in terms of morphology and brain size with Australopithecus africanus. Between 1995 and 2007, the OLAPP team has recovered teeth of eight other hominid individuals from various parts of Olduvai Gorge. These have been identified as belonging to H. habilis, Paranthropus boisei, and Australopithecus cf. africanus.     

Analysis of orientation patterns in Olduvai Bed I assemblages using GIS techniques: implications for site formation processes

Mary Leakey’s excavations at Olduvai Beds I and II provided an unparalleled wealth of data on the archaeology of the early Pleistocene. We have been able to obtain axial orientations of the Bed I bone and stone tools by applying GIS methods to the site plans contained in the Olduvai Volume 3 monograph (Leakey, 1971). Our analysis indicates that the Bed I assemblages show preferred orientations, probably caused by natural agents such as water disturbance. These results, based on new GIS techniques applied to paleoanthropological studies, have important implications for the understanding of the formative agents of Olduvai sites and the behavioral meaning of the bone and lithic accumulations in Bed I.     

The age and stratigraphic position of Olduvai Hominid I

A short history of Olduvai Hominid I is given. On the basis of absolute and relative dating its position in the Naisiusiu Bed, part of the former Bed V, is affirmed. Some connection, on morphological grounds, with other hominids in East Africa is postulated, as well as with cultural material.     

First occurrence ofXerus cf.inauris (Rodentia,Sciruridae) at Olduvai Bed I (Lower Pleistocene,Tanzania)

PalZ - Some bonodont molars of a ground squirrel are attributed toXerus cf.inauris. Coming from Olduvai Bed I fossiliferous deposits they could represent an intermediate stage between the Laetoli...     

A note on the Olduvai clavicular fragment

Certain features of the Olduvai clavicular fragment have been examined in the light of data already existing on the form of the clavicle in primates. Although similar to that of modern man, the fragment exhibits features which suggest that its function may have differed somewhat from that of the clavicle in man. The new information correlates well with that obtained from the fragmentary Sterkfontein scapula.     

A complete diversity of fossils: Perspectives

A method allowing estimation of the total diversity of fossil organisms is described and illustrated, using the examples of the Permian ammonoids of the Pamir, Permian brachiopods of Texas, and Tithonian marine animals.     

Vegetation and plant food reconstruction of lowermost Bed II, Olduvai Gorge, using modern analogs

Vegetation and plant foods for hominins of lowermost Bed II, Olduvai Gorge were modeled by examining vegetation in modern habitats in northern Tanzania (Lake Manyara, Ngorongoro, Serengeti) that are analogous to the paleolandscape in terms of climate, land forms, and soil types, as indicated by previous paleoenvironmental studies of Olduvai. Plant species in the modern habitats were identified in a series of sample plots, and those known to be eaten by modern humans, chimpanzees, or baboons were considered potentially edible for early hominins. Within the 50-80 kyr deposition of lowermost Bed II, periods of drier climate were characterized by low lake stands and a broad eastern lacustrine plain containing a mosaic of springs, marsh, woodland, and edaphic grassland. Based on results of this study, plant food diversity in each of those habitats was relatively low, but the mosaic nature of the area meant that hominins could reach several different habitat types within short distances, with access to potential plant foods including marsh plants, grass grains, roots, shrub fruits, edible parts from palms, leafy herbaceous plants, and Acacia pods, flowers, and gum. Based on Manyara analogs, a greater variety of plant foods, such as tree fruits (e.g., Ficus, Trichilia) and the roots and fruits of shrubs (e.g., Cordia, Salvadora) would be expected further east along the rivers in the lacustrine terrace and alluvial fans. Interfluves of the alluvial fans were probably less wooded and offered relatively fewer varieties of plant foods, but there is sparse paleoenvironmental evidence for the character of Olduvai's alluvial fans, making the choice of appropriate modern analogs difficult. In the western side of the basin, based on modern analogs in the Serengeti, riverine habitats provided the greatest variety of edible plant food species (e.g., Acacia, Grewia, Justicia). If the interfluves were grassland, then a large variety of potentially edible grasses and forbs were present seasonally. Periods of wetter climate resulted in a much expanded paleolake and a shrinking of the eastern lacustrine plain mosaic into a narrow zone. The dominant landscape features were then forest-lined rivers in the eastern alluvial fans, and rivers in the western side of the paleo-Olduvai basin were also better watered at these times, supporting denser woody vegetation with large varieties of edible fruits, leaves, and underground parts.     

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.     

Stable isotopes of pedogenic carbonates as indicators of paleoecology in the Plio-Pleistocene (upper Bed I), western margin of the Olduvai Basin, Tanzania

Paleosol carbonates from trenches excavated as part of a landscape-scale project in Bed I of Olduvai Gorge, Tanzania, were analyzed for stable carbon and oxygen isotopic composition. The approximately 60,000-year interval ( approximately 1.845-1.785 Ma) above Tuff IB records evidence for lake and fluvial sequences, volcanic eruptions, eolian and pedogenic processes, and the development of a fluvial plain in the western margin of the basin. Significant temporal variation in the carbonate delta(18)O values records variation of local precipitation and supports the shifts in climatic conditions interpreted from the lithologic record. During this period, carbonate delta(13)C values varied between depositional facies indicating that the paleolandscape supported a local biomass of about 40-60% C(4) plants within a mosaic of grassy woodlands and wooded grasslands. The lithologic and stable isotope record in this small lake basin indicates the area was much wetter, with more woody C(3) plants, during this interval than is the semi-arid area today. The record also reflects the variation in climatic conditions (wet/dry) documented by other global climate proxies for this time.     

The prebiotic synthesis of carbohydrates: A reassessment

Summary Questions concerning the significance of previous work on the formose reaction have led us to reexamine the question of the prebiotic synthesis of sugars. The results of new experiments lead to the following conclusions: The formose reaction is a geochemically plausible reaction which depends on neither basic conditions nor on the presence of trace amounts of carbohydrate impurities. However, this process is not a plausible source of ribose nor of any other individual sugar. In contrast to the nonspecific formation of complex mixtures of sugars via the formose reaction, the reduced sugar pentaerythritol is formed with great selectivity by the ultraviolet irradiation of 0.1 M formaldehyde. This compound may have played an important role in chemical evolution.Offprint requests to: A.W. Schwartz     

Glutathione peroxidase activity in insects: A reassessment

In vertebrate species, cytotoxic H₂O₂ and other lipid or organic hydroperoxides (ROOH) formed in aerobic metabolism are removed by a selenoprotein, glutathione peroxidase (GPOX). The GPOX activity in most rat tissues ranges from 100 to 1,000 units (1 unit = 1 nmol NADPH oxidized·mg protein^?1·min^?1), except for muscles (20–30 units). In contrast, GPOX activities of two strains of the housefly (Musca domestica), cabbage looper (Trichoplusia ni), southern armyworm (Spodoptera eridania), and black swallowtail butterfly (Papilio polyxenes), were found to be in the range 2–12 units. Trivial GPOX activity was detected in the confused flour beetle (Tribolium confusum). In the earthworm (Lumbricus terrestris), banana slug (Ariolimax columbianus), and market squid (Loligo opalescens), the GPOX activity ranged from 1 to 5 units. Tissue selenium concentrations were about 500–1,000 ppb for adult M. domestica, 600 ppb in T. confusum, 32 ppb in T. ni, 17 ppb in S. eridania, and 31 ppb in P. polyxenes larvae. The form of selenium incorporated at such high levels in tissues of invertebrates such as M. domestica remains an unresolved issue. Peroxidase activity of non-selenium glutathione-S-transferase (GT) against ROOH may compensate for the low GPOX activity. Catalase (CAT) has high activity and wide subcellular distribution in insects. This may be an evolutionary adaptation to GT's inability to catalyze the reduction of H₂O₂. The GT's peroxidase and CAT activities were not assessed for other invertebrate species, and warrants an investigation due to their reported low GPOX levels.     

Olduvai Hominin 8 foot pathology: A comparative study attempting a differential diagnosis

Olduvai Hominin (OH) 8, a 1.76 million year old left foot skeleton, has osteophytic lipping on the metatarsal bases, which when compared to a modern sample, may help paleoanthropologists determine whether the foot bones represent an injured subadult or an osteoarthritic adult. This study compares the OH 8 lipping pattern to those of 140 individual Amerindians comprising four different age classes to determine whether the OH 8 lipping is likely to be age-related osteoarthritis. OH 8 metatarsal lipping followed a pattern similar to that determined in the comparative sample to be age-related osteoarthritis. Similarities include metatarsal base lipping that is frequently located on the dorsal surface, metatarsal base lipping that is more severe on the lateral metatarsals compared to the medial metatarsals, and the presence of a pseudojoint between metatarsal 1 and metatarsal 2. The chance of finding an individual with osteoarthritis lipping increases from 3.45% in the age group 18–22 years to 55% in individuals over 35 years. The chance of finding a pseudojoint increases from 1.32% in non-osteoarthritic individuals to 15.15% in individuals with osteoarthritis. Results from this study indicate that the OH 8 foot bones are most likely from an adult and more likely to belong to Paranthropus boisei, the skull of which was found in the same excavations with OH 8, than to the juvenile Homo habilis holotype.     

First Partial Skeleton of a 1.34-Million-Year-Old Paranthropus boisei from Bed II,Olduvai Gorge,Tanzania

Recent excavations in Level 4 at BK (Bed II, Olduvai Gorge, Tanzania) have yielded nine hominin teeth, a distal humerus fragment, a proximal radius with much of its shaft, a femur shaft, and a tibia shaft fragment (cataloged collectively as OH 80). Those elements identified more specifically than to simply Hominidae gen. et sp. indet are attributed to Paranthropus boisei. Before this study, incontrovertible P. boisei partial skeletons, for which postcranial remains occurred in association with taxonomically diagnostic craniodental remains, were unknown. Thus, OH 80 stands as the first unambiguous, dentally associated Paranthropus partial skeleton from East Africa. The morphology and size of its constituent parts suggest that the fossils derived from an extremely robust individual who, at 1.338±0.024 Ma (1 sigma), represents one of the most recent occurrences of Paranthropus before its extinction in East Africa.     

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.     

Crocodylian and mammalian carnivore feeding traces on hominid fossils from FLK 22 and FLK NN 3, Plio-Pleistocene, Olduvai Gorge, Tanzania

Taphonomic analysis of the Olduvai Hominid (OH) 8 left foot from FLK NN Level 3 and the OH 35 left leg from FLK Level 22 (Zinjanthropus level) in Middle Bed I, Olduvai Gorge, indicates that both were fed upon by crocodiles. Both bear extensive tooth marking, including bisected tooth marks diagnostic of crocodylian feeding. The location of the bisected tooth marks on the distal tibia and the talus indicates disarticulation of the foot by crocodiles. The broken proximal ends of the tibia and fibula are more typical of feeding by a leopard-like carnivore, as is damage to the OH 7 mandible and parietals that are associated with and may derive from the same individual as OH 8. Previous work showing a close articulation of the foot and the leg has been used to suggest that the two specimens belong to the same individual despite deriving from sites separated by 200 m and slightly different stratigraphic levels according to previous work. The location and agent of tooth marking and the nature of gross damage do not refute this hypothesis, but the punctures on the talus and distal tibia differ in size and sharpness. Recent work shows that the stratigraphic discrepancy between OH 8 and OH 35 is greater than previously thought, refuting the single-individual hypothesis. Although seemingly unlikely, this denotes that two hominids represented by rarely found leg and foot elements both lost their left foot to crocodiles at nearby sites within a 6,000 year interval. We cannot determine if the hominids were preyed upon by crocodiles or mammalian carnivores. However, the carnivore damage to them and associated faunal remains suggests that high predation risk constrained hominid activities involving discard of the stone artifacts found at these sites. This finding is inconsistent with the interpretation of the sites as home bases or living floors.     

Examining time trends in the Oldowan technology at Beds I and II,Olduvai Gorge

The lithic analysis of the Bed I and II assemblages from Olduvai Gorge reveals both static and dynamic time trends in early hominids' technology from 1.8 to 1.2 m.y.a. The Bed I Oldowan (1.87-1.75 m.y.a.) is characterized by the least effort strategy in terms of raw material exploitation and tool production. The inclusion of new raw material, chert, for toolmaking in the following Developed Oldowan A (DOA, 1.65-1.53 m.y.a.) facilitated more distinctive and variable flaking strategies depending on the kind of raw materials. The unique characters of DOA are explainable by this raw material factor, rather than technological development of hominids. The disappearance of chert in the subsequent Developed Oldowan B and Acheulian (1.53-1.2 m.y.a.) necessitated a shift in tool production strategy more similar to that of Bed I Oldowan than DOA. However, the evidence suggests that Bed II hominids might have been more skillful toolmakers, intensive tool-users, and engaged in more active transport of stone tools than the Bed I predecessors. Koobi Fora hominids maintained a more static tool-using behavior than their Olduvai counterparts due mainly to a stable supply of raw materials. They differed from Olduvai hominids in terms of less battering of cores, consistent transport behavior, and few productions of side-struck flakes, indicating a regional variation of toolmaking and using practice. However, they shared with Olduvai hominids a temporal trend toward the production of larger flakes from larger cores after 1.6 m.y.a. Increased intake of animal resources and the expansion of ranging area of Homo ergaster would have led to the development of technological organization. Technological changes in the Oldowan industry are attested at Olduvai Gorge, Koobi Fora, and Sterkfontein, suggesting that it was a pan-African synchronous phenomenon, beginning at 1.5 m.y.a.     

Carbonate horizons, paleosols, and lake flooding cycles: Beds I and II of Olduvai Gorge, Tanzania

This study documents the petrology and stable isotope geochemistry of carbonates from six horizons from Beds I and II of Olduvai Gorge, Tanzania. The studied succession, immediately below and above Tuff IF, consists of interbedded waxy and earthy claystones with discrete carbonate horizons and thin sandstones. The succession was deposited in response to repeated flooding and withdrawal of a saline-alkaline lake. The carbonates and their overlying disconformities are important because they help define the surfaces on which hominin activity took place and allow very high-resolution correlation of geographically separated levels of hominin exploitation.The range of different carbonates includes unambiguous land-surface and pedogenic features including calcified rootmat horizons, rhizocretions, and micritic nodules, together with less determinate sparry calcite nodules. Stellate nodules are interpreted as pseudomorphs after sulfate-roses. The carbonate nodules are synsedimentary features, truncated by fluvial and other erosional surfaces. The isotopic composition of the carbonates is variable with δ¹⁸O ranging from −7.0‰ to −4.3‰, and δ¹³C from −8.5‰ to −1.6‰. A covariant increase in δ¹³C and δ¹⁸O repeats in each carbonate horizon and in individual nodules (inner to outer layers): it reflects the evolution of synsedimentary groundwaters. At times of low lake level, the carbonates started to precipitate from meteoric waters with low isotopic values and continued to form as lake levels rose and the waters became increasingly saline. Some of the samples have a last-stage cement of strontium rich dolomite, which supports late-stage flooding by the saline-alkaline lake. Previous studies of carbonate horizons from Olduvai have interpreted carbon isotope values in terms of changes in C₃ and C₄ plants that colonized the land surface. This study demonstrates that in some instances the isotope values from carbonates deposited in these lake marginal settings reflect changes in hydrology rather than vegetation.