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.     

The Neanderthal lower arm

Neanderthal forearms have been described as being very powerful. Different individual features in the lower arm bones have been described to distinguish Neanderthals from modern humans. In this study, the overall morphology of the radius and ulna is considered, and morphological differences among Neanderthals, Upper Paleolithic Homo sapiens and recent H. sapiens are described.Comparisons among populations were made using a combination of 3D geometric morphometrics and standard multivariate methods. Comparative material included all available complete radii and ulnae from Neanderthals, early H. sapiens and archaeological and recent human populations, representing a wide geographical and lifestyle range.There are few differences among the populations when features are considered individually. Neanderthals and early H. sapiens fell within the range of modern human variation. When the suite of measurements and shapes were analyzed, differences and similarities became apparent. The Neanderthal radius is more laterally curved, has a more medially placed radial tuberosity, a longer radial neck, a more antero-posteriorly ovoid head and a well-developed proximal interosseous crest. The Neanderthal ulna has a more anterior facing trochlear notch, a lower M. brachialis insertion, larger relative mid-shaft size and a more medio-lateral and antero-posterior sinusoidal shaft. The Neanderthal lower arm morphology reflects a strong cold-adapted short forearm. The forearms of H. sapiens are less powerful in pronation and supination. Many differences between Neanderthals and H. sapiens can be explained as a secondary consequence of the hyper-polar body proportions of the Neanderthals, but also as retentions of the primitive condition of other hominoids.     

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.     

The Middle-to-Upper Palaeolithic transition in Cova Gran (Catalunya, Spain) and the extinction of Neanderthals in the Iberian Peninsula

The excavations carried out in Cova Gran de Santa Linya (Southeastern PrePyrenees, Catalunya, Spain) have unearthed a new archaeological sequence attributable to the Middle Palaeoloithic/Upper Palaeolithic (MP/UP) transition. This article presents data on the stratigraphy, archaeology, and ¹⁴C AMS dates of three Early Upper Palaeolithic and four Late Middle Palaeolithic levels excavated in Cova Gran. All these archaeological levels fall within the 34-32 ka time span, the temporal frame in which major events of Neanderthal extinction took place. The earliest Early Upper Palaeolithic (497D) and the latest Middle Palaeolithic (S1B) levels in Cova Gran are separated by a sterile gap and permit pinpointing the time period in which the Mousterian disappeared from Northeastern Spain. Technological differences between the Early Upper Palaeolithic and Late Middle Palaeolithic industries in Cova Gran support a cultural rupture between the two periods. A series of 12 ¹⁴C AMS dates prompts reflections on the validity of reconstructions based on radiocarbon data. Thus, results from excavations in Cova Gran lead us to discuss the scenarios relating the MP/UP transition in the Iberian Peninsula, a region considered a refuge of late Neanderthal populations.     

Gamma-ray spectrometric dating of late Homo erectus skulls from Ngandong and Sambungmacan, Central Java, Indonesia

Hominid fossils from Ngandong and Sambungmacan, Central Java, Indonesia, are considered to be the most anatomically derived and youngest representatives of Homo erectus. Nondestructive gamma-ray spectrometric dating of three of these Homo erectus skulls showed that all samples underwent uranium leaching. Nevertheless, we could establish minimum age estimates of around 40ka, with an upper age limit of around 60 to 70ka. This means that the Homo erectus of Java very likely survived the Toba eruption and may have been contemporaneous with the earliest Homo sapiens in Southeast Asia and Australasia.     

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.     

The Liang Bua faunal remains: a 95 k.yr. sequence from Flores, East Indonesia

Excavations at Liang Bua, a limestone cave on the island of Flores, East Indonesia, have yielded a well-dated archaeological and faunal sequence spanning the last 95 k.yr., major climatic fluctuations, and two human species - H. floresiensis from 95 to 17 k.yr.¹, and modern humans from 11 k.yr. to the present. The faunal assemblage comprises well-preserved mammal, bird, reptile and mollusc remains, including examples of island gigantism in small mammals and the dwarfing of large taxa. Together with evidence from Early-Middle Pleistocene sites in the Soa Basin, it confirms the long-term isolation, impoverishment, and phylogenetic continuity of the Flores faunal community. The accumulation of Stegodon and Komodo dragon remains at the site in the Pleistocene is attributed to Homo floresiensis, while predatory birds, including an extinct species of owl, were largely responsible for the accumulation of the small vertebrates. The disappearance from the sequence of the two large-bodied, endemic mammals, Stegodon florensis insularis and Homo floresiensis, was associated with a volcanic eruption at 17 ka and precedes the earliest evidence for modern humans, who initiated use of mollusc and shell working, and began to introduce a range of exotic animals to the island. Faunal introductions during the Holocene included the Sulawesi warty pig (Sus celebensis) at about 7 ka, followed by the Eurasian pig (Sus scrofa), Long-tailed macaque, Javanese porcupine, and Masked palm civet at about 4 ka, and cattle, deer, and horse - possibly by the Portuguese within historic times. The Holocene sequence at the site also documents local faunal extinctions - a result of accelerating human population growth, habitat loss, and over-exploitation.     

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.     

Juvenile ornithopod (Dinosauria: Rhabdodontidae) remains from the Upper Cretaceous (Lower Campanian, Gosau Group) of Muthmannsdorf (Lower Austria)

The fragmentary remains of a juvenile rhabdodontid ornithopod from the Coal-bearing Complex of the Gosau Group (Lower Campanian, Grünbach syncline) at Muthmannsdorf near Wiener Neustadt, Lower Austria are revised. The material, probably belonging to a single individual, includes a right dentary (lectotype of Iguanodon suessi Bunzel, 1871, designated herein), teeth, a fragmentary parietal, fragments of scapula, ?radius, femur, tibia, two vertebrae (lost) and a manual ungual.The lectotype dentary does not provide clear autapomorphies or sufficient diagnostic features to determine its position within the Rhabdodontidae at generic level. By this “Iguanodon suessi” Bunzel, 1871 and the genus “Mochlodon” Seeley, 1881, to which it was latter referred as type species, cannot be characterized sufficiently by differential diagnosis and these are best considered nomina dubia. Based upon combined character comparisons (mainly postcranial features) the Muthmannsdorf ornithopod is referred herein to Zalmoxes Weishampel, Jianu, Csiki and Norman, 2003, a genus so far known from the late Maastrichtian of Romania. It probably but not evidently represents a yet unnamed species, most closely related to Zalmoxes shqiperorum Weishampel, Jianu, Csiki and Norman, 2003. At the present state of knowledge the Austrian material is not further diagnostic at the species level and kept in open nomenclature as Zalmoxes sp.     

New xylological data and the biogeography of the Iberian Peninsula during the Early Cretaceous

The position of the Iberian Peninsula during the Early Cretaceous, sandwiched between Laurasia and Gondwana, makes it an important area for the biogeography study of terrestrial ecosystems. Relevant data are, however, scarce. The discovery of silicified wood in the west of the Cameros Basin, in the village Hacinas (Spain), is the first record of the genus Protopodocarpoxylon in Spain and the seventh of Agathoxylon. A new species, Protopodocarpoxylon haciniensis sp. nov., is described. The anatomy of the samples studied shows that paleoclimatic conditions during the Early Cretaceous in the western part of the Cameros Basin were favorable to tree growth, with good water supply during the growth season. Although much impoverished in comparison with Western Europe, the Iberian Early Cretaceous wood floras are clearly Laurasian in affinity.     

Preface: research at Liang Bua, Flores, Indonesia

Excavations at Liang Bua, Flores, Indonesia, have yielded evidence for an endemic human species, Homo floresiensis, a population that occupied the cave between ∼95-17 ka. This discovery has major implications for early hominin evolution and dispersal in Africa and Asia, attracting worldwide interest. This preface describes the rationale for the excavations in historical, geographical, and wider research contexts, as well as the methods used. It also introduces the other papers on aspects of Liang Bua research that feature in this edition of the Journal of Human Evolution.     

The earliest European badger (Meles meles), from the Late Villafranchian site of Fuente Nueva 3 (Orce, Granada, SE Iberian Peninsula)

Paleontological and molecular data suggest that the divergence of the European and Asian badgers (Meles meles and Meles leucurus, respectively) from their ancestor Meles thorali might have taken place in the Middle to Late Villafranchian boundary (ca. 1.8 Ma). However, the available record of Late Villafrachian European badgers is scanty and poorly known, and this hypothesis needs more paleontological data in order to be tested. The badger hemimandible from the Iberian locality of Fuente Nueva 3, a locality placed in the beginning of the Late Villafranchian, supports the idea of an early divergence between European and Asian badgers.     

Distinguishing humans from great apes with AluYb8 repeats

Humans and chimpanzees share some 99% of DNA and amino acid identity, yet they exhibit important biomedical, morphological, and cognitive differences, difficult to accommodate within the remaining 1% of sequence diversity. Other types of genetic variation must be responsible for the taxonomic differences. Here we trace the evolution of AluYb8 repeats from a single origin at the roots of higher primates to a large increase in their number in humans. We identify nine AluYb8 DNA repeats in the chimpanzee genome compared to over 2200 repeats in the human, which represents a 250-fold increase in the rate of change in the human lineage and far outweighs the 99% sequence similarity between the two species. It is estimated that the average age of the human Yb8Alus is about 3.3 million years (My); almost 10% of them are identical in sequence, and hence are of recent origin. Genomic variations of this magnitude, distinguishing humans from great apes have not been realized. This explosive Alu expansion must have had a profound effect on the organization of our genome and the architecture of our chromosomes, inferentially altering profiles of gene expression and chromosome choreography in cell division. Additionally, we conclude that this major evolutionary process of Alu proliferation is driven by internal forces, written in the chemistry of DNA, rather than by external selection.     

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.