Late Pliocene hominid knapping skills: the case of Lokalalei 2C, West Turkana, Kenya

Relatively few remains of Late Pliocene hominids' knapping activities have been recovered to date, and these have seldom been studied in terms of manual dexterity and technical achievements. With regard to early hominid technological development, the evidence provided by the data from 2.34 Myr site of Lokalalei 2C (Kenya) questions both the prior assumption of a continuous and linear evolutionary trend in lithic production and the idea that it long remained static. The level of elaboration evinced by the lithic assemblage is quite unexpected in view of its age, and seemingly more advanced that what can be surmised for other Late Pliocene East-African sites, including the nearby site of Lokalalei 1. Analysis relies mainly on the dynamic reconstruction of entire cobble reduction sequences from particularly informative refitting groups. The Lokalalei 2C knappers had already internalised the notion of planning and foresight in raw material procurement and management. Beyond simple mastery of the basic technical constraints peculiar to stone knapping, they conducted a highly controlled debitage of flakes following constant technical rules and resulting in high productivity. The data suggest that early hominids displayed distinct technical competencies and techno-economic patterns of behavior, thus pointing to an intrasite complexity and intersite diversity which are not accounted for by the existing chrono-cultural classifications.     

Variation in enamel thickness within the genus Homo

Recent humans and their fossil relatives are classified as having thick molar enamel, one of very few dental traits that distinguish hominins from living African apes. However, little is known about enamel thickness in the earliest members of the genus Homo, and recent studies of later Homo report considerable intra- and inter-specific variation. In order to assess taxonomic, geographic, and temporal trends in enamel thickness, we applied micro-computed tomographic imaging to 150 fossil Homo teeth spanning two million years. Early Homo postcanine teeth from Africa and Asia show highly variable average and relative enamel thickness (AET and RET) values. Three molars from South Africa exceed Homo AET and RET ranges, resembling the hyper thick Paranthropus condition. Most later Homo groups (archaic European and north African Homo, and fossil and recent Homo sapiens) possess absolutely and relatively thick enamel across the entire dentition. In contrast, Neanderthals show relatively thin enamel in their incisors, canines, premolars, and molars, although incisor AET values are similar to H. sapiens. Comparisons of recent and fossil H. sapiens reveal that dental size reduction has led to a disproportionate decrease in coronal dentine compared with enamel (although both are reduced), leading to relatively thicker enamel in recent humans. General characterizations of hominins as having ‘thick enamel’ thus oversimplify a surprisingly variable craniodental trait with limited taxonomic utility within a genus. Moreover, estimates of dental attrition rates employed in paleodemographic reconstruction may be biased when this variation is not considered. Additional research is necessary to reconstruct hominin dietary ecology since thick enamel is not a prerequisite for hard-object feeding, and it is present in most later Homo species despite advances in technology and food processing.     

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.     

Correlation of the KHS Tuff of the Kibish Formation to volcanic ash layers at other sites, and the age of early Homo sapiens (Omo I and Omo II)

Hominin specimens Omo I and Omo II from Member I of the Kibish Formation, Ethiopia are attributed to early Homo sapiens, and an age near 196 ka has been suggested for them. The KHS Tuff, within Member II of the Kibish Formation has not been directly dated at the site, but it is believed to have been deposited at or near the time of formation of sapropel S6 in the Mediterranean Sea. Electron microprobe analyses suggest that the KHS Tuff correlates with the WAVT (Waidedo Vitric Tuff) at Herto, Gona, and Konso (sample TA-55), and with Unit D at Kulkuletti in the Ethiopian Rift Valley. Konso sample TA-55 is older than 154 ka, and Unit D at Kulkuletti is dated at 183 ka. These correlations and ages provide strong support for the age originally suggested for the hominin remains Omo I and Omo II, and for correlation of times of deposition in the Kibish region with formation of sapropels in the Mediterranean Sea. The Aliyo Tuff in Member III of the Kibish Formation is dated at 104 ka, and correlates with Gademotta Unit 15 in the Ethiopian Rift Valley.     

Hallucial convergence in early hominids

There is a richly documented fossil record of the evolutionary transition from ape-sized brains that are less that one-third the size of modern humans through a series of intermediate-sized brains up to the modern range. The first report on the discovery of the foot of the Stw 573 skeleton emphasized the apparent transitional nature of its great toe [Clarke, R.J., Tobias, P.V., 1995. Sterkfontein Member 2 foot bones of the oldest South African hominid. Science 269, pp. 521-524]. The hallux appeared to be intermediate in its divergence between human-like adduction and ape-like abduction. A major part of this evidence is the medial encroachment of the metatarsal I facet on the medial cuneiform. This study quantifies the variability of this feature in extant hominoids and fossil hominids. The results are consistent with the view that all currently known hominids were specialized for bipedality and lacked the ape-like ability to oppose the great toe.     

Presence of Hypolagus Dice, 1917 (Mammalia, Lagomorpha) in the Neogene of the Balearic Islands (Western Mediterranean): Description of Hypolagus balearicus nov. sp.

In this paper we describe a new species of Hypolagus coming from an early Pliocene karstic deposit near Caló den Rafelino (Manacor, Mallorca). It represents the westernmost European record for the genus. The faunal assemblage of this deposit represents an early phase of the second insular faunal episode of Mallorca, related with the Messinian regressive episode (latest Miocene). Although the postcranial are relatively robust, the size of all these elements is included in the range of continental leporids. It is unknown if they display significant allometric changes, as occurred in other insular leporids. p3 is small and displays a markedly trapezoidal outline and a deep hypoflexid (between 54 and 56% of the total tooth width). The presence of Hypolagus in the Neogene of the Balearic Islands is in agreement with the faunal scenario for the European continent during the late Miocene and the Pliocene, where this genus is abundant and widely distributed.     

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.     

Carcass transport decisions in Homo antecessor subsistence strategies

Pleistocene foragers used several prey acquisition and processing strategies. These strategies and their associated decisions are elucidated by taphonomic studies that cover animal transport, modifications by different agents and archaeological remains. Interpretative models of archaeological sites are by necessity based on natural and experimental observations. Ethno-archaeological data shows that several factors influenced decisions about carcass transport from the kill site to the home site. These factors often have little archaeological visibility. Díez et al. (1999) has previously interpreted the general characteristics of the macro-mammal remains from Gran Dolina Level TD6-2 (Sierra de Atapuerca, Burgos, Spain) as the result of anthropic accumulation, in which the anatomical profiles appeared to be the result of selective transport based on the animals’ weight. Recent taphonomic analysis has shown that carcasses with different weights may be subject to similar transport strategies, suggesting that other factors influenced these choices. The hominins that occupied TD6-2 (the TD6-2 hominin group), at least sometimes, transported large carcasses to the cave in their entirety, implying participation by groups of individuals in hunting parties. These individuals delayed their consumption of large amounts of food, instead moving it to Gran Dolina, where it was shared with other group members. These decisions are evidence of social cooperation and food sharing amongst early European hominins.     

A new species of the genusAustralopithecus (primates: hominidae) from Plio/Pleistocene deposits west of Lake Turkana in Kenya

The cranium of a robust australopithecine, KNM WT 17000, was discovered from the Plio/Pleistocene deposits west of Lake Turkana in Kenya, and assigned to the speciesAustralopithecus boisei Leakey, 1959. A comparative morphological study shows that it does not conform with the diagnosis forA. boisei. It is characterized by having a much smaller brain, a low hyperprognathous facial skeleton, and a less developed masticatory apparatus. Its unique morphological pattern justifies its placement in a new taxon which is calledAustralopithecus walkeri n. sp.     

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.     

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.     

A new skeleton of Theropithecus brumpti (Primates: Cercopithecidae) from Lomekwi, West Turkana, Kenya

A relatively complete skeleton of the fossil papionin, Theropithecus brumpti, from the site of Lomekwi, west of Lake Turkana, Kenya, is here described. The specimen, KNM-WT 39368, was recovered at the site of LO 5 (3°51′N and 35°45′E), from sediments dated to approximately 3·3 Ma. The skeleton is that of an old adult male and preserves a number of articulated elements, including most of the forelimbs and tail. The cranial morphology is that of a large, early T. brumpti, exhibiting a deep mandible with a deeply excavated mandibular corpus fossa, and mandibular alveoli and cheek teeth arrayed in a reversed Curve of Spee. The forelimb skeleton exhibits a unique mixture of characteristics generally associated with a terrestrial locomotor habitus, such as a narrow scapula and a highly stable elbow joint, combined with those more representative of habitual arborealists, such as muscle attachments reflecting a large rotator cuff musculature and a flexible shoulder joint. The forelimb of KNM-WT 39368 also presents several features, unique toTheropithecus , which represent adaptations for manual grasping and fine manipulation. These features include a large, retroflexed medial humeral epicondyle (to which large pronator, and carpal and digital flexor muscles attached) and proportions of the digital rays that denote capabilities for precise opposition between the thumb and index finger. Taken together, these features indicate that one of the earliest recognized representatives of Theropithecus exhibited the food harvesting and processing anatomy that distinguished the genus through time and that contributed to its success throughout the later Pliocene and Pleistocene. Based on the anatomy of KNM-WT 39368 and the known habitat preference of T. brumpti, the species is reconstructed as being a generally terrestrial but highly dexterous, very large-bodied, sexually dimorphic, and possibly folivorous papionin. T. brumpti was adapted for propulsive quadrupedal locomotion over generally even ground, and yet was highly adept at manual foraging. The estimate of 43·8 kg body mass for KNM-WT 39368 renders unlikely the possibility that the species, or at least adult males of the species, were highly arboreal. T. brumpti, as represented by KNM-WT 39368, is seen as a large, colorfully decorated, and basically terrestrial papionin that was restricted to riverine forest habitats in the Lake Turkana Basin from the middle to latest Pliocene.     

First record of a nematode Metastrongyloidea (Aelurostrongylus abstrusus larvae) in Achatina (Lissachatina) fulica (Mollusca, Achatinidae) in Brazil

Achatina (Lissachatina) fulica was introduced in Brazil in the 1980s for commercial purposes (“escargot” farming) and nowadays, mainly by human activity, it is widespread in at least 23 out of 26 Brazilian states and Brasília, including the Amazonian region and natural reserves, where besides a general nuisance for people it is a pest and also a public health concern, since it is one of the natural intermediate host of Angiostrongylus cantonensis, ethiological agent of the meningoencephalitis in Asia. As Brazil is experiencing the explosive phase of the invasion, the Laboratório de Malacologia do Instituto Oswaldo Cruz/Fiocruz has been receiving samples of these molluscs for identification and search for Angiostrongylus cantonensis and Angiostrongylus costaricensis larvae. While examining samples of A. fulica different nematode larvae were obtained, including Aelurostrongylus, whose different species are parasites of felids, dogs, primates, and badger. Morphological and morphometric analyses presented herein indicated the species Aelurostrongylus abstrusus, as well as the occurrence of other nematode larvae (Strongyluris-like) found in the interior of the pallial cavity of A. fulica. This is the first report in Brazil of the development of A. abstrusus infective larvae in A. fulica evidencing the veterinary importance of this mollusc in the transmission of A. abstrusus to domestic cats. Since the spread of A. fulica is pointed out in the literature as one of the main causative spread of the meningoencephalitis caused by A. cantonensis the authors emphasize the need of sanitary vigilance of snails and rats from vulnerable areas for A. cantonensis introduction as the port side areas.     

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.