Taxonomic affinities and evolutionary history of the Early Pleistocene hominids of Java: dentognathic evidence

Temporal changes, within-group variation, and phylogenetic positions of the Early Pleistocene Javanese hominids remain unclear. Recent debate focused on the age of the oldest Javanese hominids, but the argument so far includes little morphological basis for the fossils. To approach these questions, we analyzed a comprehensive dentognathic sample from Sangiran, which includes most of the existing hominid mandibles and teeth from the Early Pleistocene of Java. The sample was divided into chronologically younger and older groups. We examined morphological differences between these chronological groups, and investigated their affinities with other hominid groups from Africa and Eurasia. The results indicated that 1) there are remarkable morphological differences between the chronologically younger and older groups of Java, 2) the chronologically younger group is morphologically advanced, showing a similar degree of dentognathic reduction to that of Middle Pleistocene Chinese H. erectus, and 3) the chronologically older group exhibits some features that are equally primitive as or more primitive than early H. erectus of Africa. These findings suggest that the evolutionary history of early Javanese H. erectus was more dynamic than previously thought. Coupled with recent discoveries of the earliest form of H. erectus from Dmanisi, Georgia, the primitive aspects of the oldest Javanese hominid remains suggest that hominid groups prior to the grade of ca. 1.8-1.5 Ma African early H. erectus dispersed into eastern Eurasia during the earlier Early Pleistocene, although the age of the Javanese hominids themselves is yet to be resolved. Subsequent periods of the Early Pleistocene witnessed remarkable changes in the Javanese hominid record, which are ascribed either to significant in situ evolution or replacement of populations.     

Hominid cranial remains from upper Pleistocene deposits at Aduma, Middle Awash, Ethiopia

The Upper Pleistocene localities of Aduma and Bouri have yielded hominid fossils and extensive Middle Stone Age (MSA) archaeological assemblages. The vertebrate fossils recovered include parts of four hominid crania from Aduma and a complete right parietal from Bouri. Archaeological associations and radiometric techniques suggest an Upper Pleistocene age for these hominids. The more complete cranium from Aduma (ADU-VP-1/3) comprises most of the parietals, the occipital, and part of the frontal. This cranium is compared to late Middle and Upper Pleistocene hominid crania from Africa and the Middle East. The Aduma cranium shows a mosaic of cranial features shared with "premodern" and anatomically modern Homo sapiens. However, the posterior and lateral cranial dimensions, and most of its anatomy, are centered among modern humans and resemble specimens from Omo, Skhul, and Qafzeh. As a result, the Aduma and Bouri Upper Pleistocene hominids are assigned to anatomically modern Homo sapiens.     

Endocranial capacity of the bodo cranium determined from three-dimensional computed tomography

The 600,000-year-old cranium from Bodo, Ethiopia, is the oldest and most complete early Middle Pleistocene hominid skull from Africa. "Virtual endocast" models created by three-dimensional computed tomography (CT) techniques indicate an endocranial capacity of about 1,250 cc for this cranium (with a reasonable range between approximately 1,200-1,325 cc, depending on how missing portions of the basicranial region are reconstructed). From these determinations, several important implications emerge concerning current interpretations of "tempo and mode" in early hominid brain evolution: 1) already by the early Middle Pleistocene, at least one African hominid species, Homo heidelbergensis, had reached an endocranial capacity within the normal range of modern humans; 2) in spite of its large endocranial capacity, estimates of Bodo's encephalization quotient fall below those found in a large sample of Homo sapiens (both fossil and recent) and Neandertals; and 3) the greatest burst of brain expansion in the Homo lineage may not have been in the last several hundred thousand years, but rather much earlier in the Lower to early Middle Pleistocene.     

A new hominid parietal from Bodo, Middle Awash Valley, Ethiopia

A piece of left parietal of a Middle Pleistocene hominid, recovered from the Upper Bodo Sand Unit, in the Middle Awash, Ethiopia, is described anatomically and compared to Middle Pleistocene hominids and modern Homo sapiens. It bears several primitive features and has important implications for the original Bodo skull, found at the same stratigraphic level in the same area. The new fossil skull represents a different individual from the original Bodo skull.     

A review of the chronology of the European Middle Pleistocene hominid record

Over the last seventy years, European hominid fossils and associated archaeological remains have been dated by reference to the classical, fourfold glacial/interglacial subdivision of the Pleistocene. This method seemed relatively straightforward and precise especially as new discoveries were fitted into the schemes, apparently strengthening the correlations and increasing their validity. However, recent studies of deep-sea cores and terrestrial deposits, combined with new developments in relative and absolute dating, have shown that the fourfold schemes are oversimplified. This paper critically reviews some of the dating evidence from sites with hominid remains generally considered as Middle Pleistocene (ca. 700,000–128,000 BP). The hominid and archaeological remains are shown to require independent dating and a cautious approach is adopted towards the use of mammalian faunal remains as chronological indicators. The techniques and results of absolute dating are discussed with reference to present problems and future prospects. At a time of transition from an old framework to new correlations, it is inevitable that some conclusions seem tentative and others rather negative. Nevertheless, the Middle Pleistocene in Europe is more fully understood and better dated than the equivalent period in other parts of the world.     

Excavations at new sites of early man in Murcia: Sima de las Palomas del Cabezo Gordo and Cueva Negra del Estrecho del Río Quípar de la Encarnación

Two new sites with hominid, palaeolithic and faunal remains from the late Middle and early Upper Pleistocene in Murcia, S.E. Spain, are presented in the light of recent ongoing excavations and multidisciplinary research. The article outlines the origins and development of the research project with reference to aspects of field strategy and methodology, and comments upon the significant hominid and middle palaeolithic findings.     

The Boxgrove,England, middle Pleistocene herpetofauna: Paleogeographic,evolutionary, stratigraphic,and paleoecological relationships

Triturus vulgaris (smooth newt), Triturus helveticus or T. vulgaris (palmate or smooth newt), Triturus sp. (newt), Pelobates fuscus (common spadefoot), Bufo bufo (common toad), Bufo calamita (natterjack toad), Bufo sp. (toad), Rana arvalis (moor frog), Rana temporaria (common frog), Rana sp. (frog), Anguis fragilis (slow worm), Lacerta cf. L. vivipara (common lizard), Natrix natrix (grass snake), and Natrix sp. (grass, viperine, or dice snake) were identified at the Middle Pleistocene Boxgrove Site, West Sussex, England. This is the first British fossil record of Pelobates fuscus and the earliest fossil record in Britain for the endangered species Bufo calamita. All of these herpetological species are extant and all of them occur in Britain today with the exception of Pelobates fuscus and Rana arvalis that presently live on the European continent.

The Boxgrove, Westbury, and West Runton British pre‐Anglian Middle Pleistocene herpetofaunas show no apparent differences among themselves in patterns of species composition, diversity, or number of exotics. But these three herpetofaunas together have [1] less species diversity and [2] fewer exotic continental species than in the Cudmore Grove British post‐Anglian Middle Pleistocene herpetofauna.

Only the Terrestral Sequence Unit at Boxgrove yielded enough herpetological species for paleoecological interpretation. These taxa indicate a quiet pool surrounded by a somewhat humid vegetated area that gave way to a more xerophytic sandy area, and a paleoclimate at least as warm and perhaps somewhat warmer than occurs in the area today.     

On the age of the hominid fossils at the Sima de los Huesos, Sierra de Atapuerca, Spain: paleomagnetic evidence

We report new paleomagnetic data for the Middle Pleistocene hominid-bearing strata in the Sima de los Huesos, North Spain. Sediments (brown muds with human and bear fossils and the underlying sterile clayey and sandy unit) preserve both normal and reversed magnetic components. The sterile unit has exclusively reversed magnetization, dating back to the Matuyama Chron, and thus is Lower Pleistocene in age. The overlying fossiliferous muds have a dominant normal magnetization that overprints a partially resolved reversed magnetization. These data are compatible with one of the reversal events that occurred during the Brunhes Chron. Combined with the existing U-series dates and evidence from the macro- and microfauna, these paleomagnetic results suggest an age of the hominid fossils between 325 to 205 ka, whereas the underlying sand and silts are older than 780 ka.     

Human mandibular incisors from the late Middle Pleistocene locality of Hoedjiespunt 1, South Africa.

The Hoedjiespunt 1 locality is an archaeological and palaeontological site located on the Hoedjiespunt Peninsula at Saldanha Bay, South Africa. In 1996 two human teeth, a left central mandibular incisor and a left lateral mandibular incisor, were discovered during excavations in the late Middle Pleistocene palaeontological layers. These teeth are described and are found to belong to a single subadult individual. Despite their developmental stage, these incisors already display early signs of wear. Their crown diameters are larger than modern and archaeological African comparative material and are most closely comparable with crown diameters of an early Middle Pleistocene and late Middle Pleistocene dental sample from Africa, Europe and Asia. In the light of this metrical evidence, data on two previously excavated maxillary molars, most probably belonging to the same individual, were re-examined. It was found that the Hoedjiespunt 1 hominid possessed dental metrical features (large anterior teeth and small molars) comparable with other African and European hominids referred to the Middle Pleistocene.     

Early paleolithic of China and eastern Asia

In attempting to understand the course of human evolution and the nature of hominid adaptation over the past few million years, it is necessary to consider prevailing evidence from all parts of the world. Eastern Asia provides a range of important questions and challenges with regard to this evolutionary puzzle. Although evidence for earlier ape evolution is present in China (for example, at Lufeng in Yunnan Province), the earliest evidence for hominid presence appears to be in the Early Pleistocene, apparently the result of a migration of hominids to and subsequent adaptation within Eastern Asia. The archeological record provides a closer look at some technological aspects of this adaptation during the Early and Middle Pleistocene, showing both distinctive contrasts and intriguing continuities relative to the rest of the Old World.     

Cut marks on the Bodo cranium: a case of prehistoric defleshing

Cut marks were discovered on the Middle Pleistocene Bodo cranium from Ethiopia. The cut marks most closely resemble experimental damage caused by the application of stone tools to fresh bone. This discovery constitutes the earliest solid evidence for intentional defleshing of a human ancestor and offers new research avenues for the investigation of early hominid mortuary practices.     

The ATD6-5 mandibular specimen from Gran Dolina (Atapuerca, Spain). Morphological study and phylogenetic implications.

Metric and shape features of the Lower Pleistocene mandibular specimen ATD605 from the level 6 of Gran Dolina site (Atapuerca, Spain) are compared with a large sample of fossil hominid mandibles. The analysis shows that ATD6-5 displays a generalized morphology largely shared with both African and European Lower and Middle Pleistocene samples. However, distinctive African traits, such as corpus robustness and strong alveolar prominence, are absent in the Gran Dolina specimen. At the same time, none of the apomorphic features that characterize Middle and early Upper Pleistocene European hominids can be recognized in ATD6-5. Finally, the Gran Dolina specimen displays a remarkable position of the mylohyoid groove, only comparable to that found in immature specimens of Homo ergaster, and very rarely in adult H. sapiens. The morphology of ATD6-5 supports the hypothesis of an African origin for the first Europeans with subsequent phylogenetic continuity with Middle Pleistocene populations in Europe. These findings are consistent with H. antecessor being the last common ancestor of Neandertals and H. sapiens.     

The M. obturator internus sulcus on Middle and Late Pleistocene human ischia

Recent human ischia and those of Middle and Late Pleistocene hominids exhibit variation in the cranio-caudal location of the sulcus for the internal obturator muscle as it rounds the ischium through the lesser sciatic notch, from being fully cranial of the ischial tuberosity, to bordering the tuberosity, to crossing the superior tuberosity. Among two recent human samples, all three forms exist, with the cranial position of the sulcus being more common in a 20th century Euroamerican sample whereas the intermediate one predominates in a horticultural late prehistoric Amerindian sample. The available Pleistocene Homo fossil remains exhibit the full range of variation with no one form being dominant in Middle Pleistocene archaic humans and Middle Paleolithic late archaic and early modern humans. It is only within the Upper Paleolithic that the cranial and intermediate locations for the sulcus become predominant. These patterns therefore indicate that it is inappropriate to use this feature for distinguishing later Pleistocene hominid groups. © 1996 Wiley-Liss, Inc.     

Middle Pleistocene ecology and Neanderthal subsistence: Insights from stable isotope analyses in Payre (Ardèche,southeastern France)

The Middle Palaeolithic site of Payre in southeastern France yields abundant archaeological material associated with fossil hominid remains. With its long sequence of Middle Pleistocene deposits, Payre is a key site to study the Middle Palaeolithic chronology of this region. This study is the first to investigate carbon and oxygen isotope contents of Neanderthal tooth enamel bioapatite, together with a wide range of herbivorous and carnivorous species. The aim is to contribute to the understanding of hunting behaviour, resource partitioning, diet and habitat use of animals and Neanderthals through a palaeoecological reconstruction.     

Patterns of evolution in Middle Pleistocene hominids

This paper reviews the chronology and morphological variability of Middle Pleistocene H. erectus. specimens. Functional complexes are delineated within the skull and dentition, and their total morphological patterns quantified using univariate and multivariate statistical analysis. Statistical distances are calculated between H. erectus and other hominid samples for each complex, compared to illustrate patterns of mosaic evolution within the skull and dentition of middle Quaternary hominids, and estimated evolution rates are derived. An attempt is made to relate the observed morphological patterns to ecological shifts by early hominid communities, and to assess their significance for hominid taxonomy.     

In the Footprints of Our Ancestors: An Overview of the Hominid Track Record

Hominid footprints are particularly appealing and evocative of the living activity of our ancestors. The most famous and oldest (Late Pliocene, ca. 3.7 Ma) hominid footprints, from Laetoli in East Africa, have been attributed, with some uncertainly, to genus Homo or Australopithecus. The African track record also yields Early Pleistocene (～1.5 Ma) tracks attributable to Homo erectus. The only well-documented Middle Pleistocene tracks (age ～325,000-385,000 yrs) are reported from Italy and presumably represent a pre-Homo sapiens species.

The oldest Late Pleistocene tracks (～117,000 yrs), from southern Africa, may represent modern humans. However, the majority of Late Pleistocene sites are European, associated with caves in Romania, Greece, France and elsewhere, where hominid track preservation is often of high quality. Dates range from ～10,000 to ～62,000 BP Cavesite mammal tracks are almost exclusively those of carnivores, thus representing a distinctive underground ecology. Late Pleistocene open air sites are reported from widely scattered locations in Africa, Turkey, Tibet, Korea, Australia and even in the New World (Chile, Argentina and Mexico).

Early to Middle Holocene sites (> ～4,000 yrs BP) mainly occupy riparian, lacustrine, estuarine and littoral settings where the ichnofaunas are dominated by ungulates and shorebirds. Among these sites from England, Nicaragua, Argentina and Mexico and the United States, a few have been described in some detail. Younger Holocene sites are frequently associated with specified cultural periods (e.g., Neolithic, Bronze Age) or specific indigenous cultures, where supplemental archeological evidence may be directly associated with the footprint evidence.

At most surficial and some subterranean hominid tracksites, mammal and/or bird tracks are quite common and of use in creating a paleoecological picture of local faunas. The global distribution of human and hominid tracks is consistent with body fossil evidence and the record of archeological, cultural artifacts. However, in a few cases tracks suggest colonization of certain regions (Tibetan Plateau and the New World) earlier than previously thought. Tracks also give clues to behavior, age and health status of the trackmakers.     

Neanderthals: Names, Hypotheses, and Scientific Method

The Neanderthal "problem" is less likely to be solved by hypotheses which are broad and inclusive, with respect to late Pleistocene hominids, than by more limited ones for which existing data are suitable and which conform in as many dimensions as possible to evolutionary theory. Specifically, the Brose-Wolpoff hypothesis (1971) fails to use a "population" understanding of Upper Pleistocene hominid variation and argues a universal model of Middle Paleolithic-Upper Paleolithic cultural and physical transition unsuitable for southern Africa or Southeast Asia.     

Two hominin incisor teeth from the middle Pleistocene site of Boxgrove, Sussex, England

In 1995-1996 two isolated hominin lower incisors were found at the middle Pleistocene site of Boxgrove in England, with Lower Palaeolithic archaeology. Boxgrove 2 is a permanent lower right central incisor and Boxgrove 3 a permanent lower left lateral incisor. They were found separately, but close to one another and appear to belong to the same individual. The Boxgrove 1 tibia discovered in 1993 came from a different stratigraphic context and is thus believed to represent a different individual. This paper describes the morphology of the incisors, which is similar to other middle Pleistocene hominin specimens and, as with the tibia, suggests that they could be assigned to Homo heidelbergensis (recognising that the taxonomic status of this species is still a matter of debate). The incisors show substantial attrition associated with secondary dentine deposition in the pulp chamber and clearly represent an adult. They also show extensive patterns of non-masticatory scratches on the labial surfaces of both crown and root, including some marks which may have been made postmortem. The roots were exposed in life on their labial sides by a large dehiscence, extending almost to the root apex. This is demonstrated by deposits of calculus, polishing, and scratching on the exposed surfaces. The dehiscence may have been caused by repeated trauma to the gingivae or remodelling of the tooth-supporting tissues in response to large forces applied to the front of the dentition.     

Dispersal and colonisation, long and short chronologies: how continuous is the Early Pleistocene record for hominids outside East Africa?

This paper examines the evidence for hominids outside East Africa during the Early Pleistocene. Most attention has focused recently on the evidence for or against a late Pliocene dispersal, ca. 1.8 Ma., of hominids out of Africa into Asia and possibly southern Europe. Here, the focus is widened to include North Africa as well as southern Asia and Europe, as well as the evidence in these regions for hominids after their first putative appearance ca. 1.8 Ma. It suggests that overall there is very little evidence for hominids in most of these regions before the Middle Pleistocene. Consequently, it concludes that the colonising capabilities of Homo erectus may have been seriously over-rated, and that even if hominids did occupy parts of North Africa, southern Europe and southern Asia shortly after 2 Ma, there is little evidence of colonisation. Whilst further fieldwork will doubtless slowly fill many gaps in a poorly documented Lower Pleistocene hominid record, it appears premature to conclude that the appearance of hominids in North Africa, Europe and Asia was automatically followed by permanent settlement. Rather, current data are more consistent with the view that Lower Pleistocene hominid populations outside East Africa were often spatially and temporally discontinuous, that hominid expansion was strongly constrained by latitude, and that occupation of temperate latitudes north of latitude 40 degrees was largely confined to interglacial periods.     

Posterior dental size reduction in hominids: The Atapuerca evidence

In order to reassess previous hypotheses concerning dental size reduction of the posterior teeth during Pleistocene human evolution, current fossil dental evidence is examined. This evidence includes the large sample of hominid teeth found in recent excavations (1984–1993) in the Sima de los Huesos Middle Pleistocene cave site of the Sierra de Atapuerca (Burgos, Spain). The lower fourth premolars and molars of the Atapuerca hominids, probably older than 300 Kyr, have dimensions similar to those of modern humans. Further, these hominids share the derived state of other features of the posterior teeth with modern humans, such as a similar relative molar size and frequent absence of the hypoconulid, thus suggesting a possible case of parallelism. We believe that dietary changes allowed size reduction of the posterior teeth during the Middle Pleistocene, and the present evidence suggests that the selective pressures that operated on the size variability of these teeth were less restrictive than what is assumed by previous models of dental reduction. Thus, the causal relationship between tooth size decrease and changes in food-preparation techniques during the Pleistocene should be reconsidered. Moreover, the present evidence indicates that the differential reduction of the molars cannot be explained in terms of restriction of available growth space. The molar crown area measurements of a modern human sample were also investigated. The results of this study, as well as previous similar analyses, suggest that a decrease of the rate of cell proliferation, which affected the later-forming crown regions to a greater extent, may be the biological process responsible for the general and differential dental size reduction that occurred during human evolution. © 1995 Wiley-Liss, Inc.