The current state of korean paleoanthropology

The hominid fossil and Paleolithic archaeology records from the Korean Peninsula are extensive, but relatively little is known about the Korean human evolutionary record outside this region. The Korean paleoanthropological record is reviewed here in light of major research issues, including the hominid fossil record, relative and chronometric dating, lithic analysis, hominid subsistence, and the presence of bone tools, art and symbolism. Some of the major conclusions drawn from this review include: (1) hominid fossils have been found in nine separate sites on the Korean Peninsula; (2) possible Homo erectus fossils are present in North Korea; (3) Ryonggok Cave, in North Korea, has exposed the remains of at least five archaic Homo sapiens individuals; (4) a possible burial of an anatomically modern Homo sapiens child, discovered in Hungsu Cave in South Korea, has been tentatively dated to roughly 40,000 years ago; (5) handaxes and cleavers have been found at a number of sites near Chongokni and they appear to date to at least 100,000 years ago; and (6) taphonomic studies are necessary for addressing issues related to determining the nature of hominid-carnivore interaction over similar resources (e.g. carcasses and shelter); and the presence/absence of Early Paleolithic bone tools, art, and symbolism in Korea.     

Australopithecus afarensis and the single species hypothesis

Ferguson (1989) has recently argued that the variability seen in the fossils assigned toA. afarensis is far more than expected for a single hominid species, and therefore proposes they represent multiple taxa. In particular, he utilizes data on variation in dental metrics and in premolar morphology in support of this hypothesis. A re-evaluation of these data finds the above conclusion to be unwarranted. Variation in dental metrics providesno basis for separating this sample into multiple taxa, regardless of the analog that is used (i.e. modern primate species or fossil hominid species). Additionally, data on P₃ morphology indicate that thepattern of variation seen in the Laetoli/Hadar sample is comparable to the sexual variation seenwithin a single hominoid species. Overall, the balance of the evidence at present indicates that the fossils from Laetoli and Hadar represent a single hominid species,A. afarensis.     

Origin and development of Tool-making behavior in Africa and Asia

Around two-and-a-half million years ago, some hominid populations in Africa began to modify stones and bones in a manner that can be recognized by prehistorians as artifacts, and, by definition, produced the earliest identifiable archaeological record. It is likely that earlier hominid groups also may have had relatively rich tool-using behavioral repertoires similar to that seen in modern chimpanzees (McGrew, 1992), such tools may have been made of perishable materials or minimally modified and thus difficult to identify. This review will focus on the earliest archaeological traces and the spread of hominids out of Africa and into Eurasia.     

Hominid fossil sample from Kanjera,Kenya: Description,provenance, and implications of new and earlier discoveries

Anatomically modern hominids were first collected from Kanjera, Kenya, by L.S.B. Leakey in the 1930s. Their apparent association with an archaic fauna was quickly challenged, throwing their age into doubt. Further unpublished hominid fragments were collected in 1974, 1975, 1981, and 1987. We review the context and morphology of the entire hominid sample. A minimum number of five individuals is represented by both cranial and postcranial elements. Several individuals have thickened cranial vaults, a characteristic originally thought to reflect their great antiquity. Vault thickening resulted from diploic expansion and may have been a response to acquired or inherited anemia. The entire hominid sample postdates the Kanjera Formation, deposited from the early into the middle Pleistocene. Most of the sample was derived from the black cotton soil capping the stratigraphic column. The morphology and context of the Kanjera hominids is consistent with human skeletal remains from nearby Holocene sites. Hominid 3 was probably an intrusive burial into an early Pleistocene bed. © 1995 Wiley-Liss, Inc.     

Hominid Ichnotaxonomy: An Exploration of a Neglected Discipline

Although more than 60 ancient hominid track sites ranging in age from 3.7 million to less than 500 B. P. are recorded from all continents except Antarctica, no ichnotaxonomic names have ever been formally proposed for hominid tracks. There is no prohibition to the naming of fossil footprints of species that created tracks and trackways similar to those of living species. On the contrary, there is precedent for the naming of ichnotaxa corresponding to the dominant extant vertebrates classes: mammals = Mammalipedia and birds = Avipeda. The hominid track site sample includes only about a dozen sites where footprint preservation is good enough to show details of diagnostic foot morphology and typical trackway morphology. We infer that the Acahualinca Footprint Museum site in Nicaragua represents the most important ancient hominid track site that combines accessibility, a large sample of well-preserved trackways and reliable dating. For this reason, we select the Nicaraguan tracks as the type sample for the new ichnotaxon Hominipes modernus ichnogen., and ichnsp. et ichnosp. nov., which we infer to represent fully modern Homo sapiens. Our preliminary investigations of other track sites suggest that the majority also yield H. modernus. However, at many sites preservation is insufficient to make an ichnotaxonomic designation at the species level or to infer that the trackmaker was H. sapiens. Thus, at many sites including the famous Laetoli site, we apply the more general label of Hominipes isp. indet.     

Human taxonomic diversity in the pleistocene: Does Homo erectus represent multiple hominid species?

Recently, nomina such as “Homo heidelbergensis” and “H. ergaster” have been resurrected to refer to fossil hominids that are perceived to be specifically distinct from Homo sapiens and Homo erectus. This results in a later human fossil record that is nearly as speciose as that documenting the earlier history of the family Hominidae. However, it is agreed that there remains only one extant hominid species: H. sapiens. Has human taxonomic diversity been significantly pruned over the last few hundred millennia, or have the number of taxa been seriously overestimated? To answer this question, the following null hypothesis is tested: polytypism was established relatively early and the species H. erectus can accommodate all spatio-temporal variation from ca. 1.7 to 0.5 Ma. A disproof of this hypothesis would suggest that modern human polytypism is a very recent phenomenon and that speciation throughout the course of human evolution was the norm and not the exception. Cranial variation in a taxonomically mixed sample of fossil hominids, and in a modern human sample, is analyzed with regard to the variation present in the fossils attributed to H. erectus. The data are examined using both univariate (coefficient of variation) and multivariate (determinant) analyses. Employing randomization methodology to offset the small size and non-normal distribution of the fossil samples, the CV and determinant results reveal a pattern and degree of variation in H. erectus that most closely approximates that of the single species H. sapiens. It is therefore concluded that the null hypothesis cannot be rejected. © 1993 Wiley-Liss, Inc.     

Chin morphology and sexual dimorphism in the fossil hominid mandible sample from Klasies River Mouth

The site of Klasies River Mouth (KRM) in South Africa has produced a small sample of early Upper Pleistocene hominid remains that have been a focus for discussions of the origins of modern humans. Despite certain primitive characteristics exhibited by these fossils, proponents of a single recent origin have attributed them to early modern humans. Critics of this hypothesis have emphasized the significance of the archaic features evident in this sample, including the absence of pronounced chins among the mandibular specimens. This study compares the size range and chin morphology exhibited by the KRM mandibles with that of Neandertals, Upper Pleistocene humans, and recent humans. The extreme sexual dimorphism documented among the KRM fossils reflects the presence of a very small individual, and previous efforts to classify the KRM sample as archaic on the basis of their robusticity have failed to address the significance of this diminutive hominid. While each KRM fossil falls within the 95% envelope of variability established for chin development in a comparative modern sample, a similarly low frequency of pronounced chins is very unlikely to be found in any recent human population. The morphological pattern of the KRM mandibles is clearly distinct from that of Neandertals and of recent humans. © 1996 Wiley-Liss, Inc.     

An archaic character in the Broken Hill innominate E. 719

The additional hominid material from Broken Hill, Kabwe, Zambia, is only dubiously associated with the hominid cranium from the site and is often considered to be anatomically modern in morphology. This study identifies an archaic feature, previously recognised in Pliocene and earlier Pleistocene innominates, in the Broken Hill innominate E. 719. An acetabulo-cristal buttress of cortical bone 10 mm thick is present, and this can be clearly distinguished from the morphology present in a comparative sample of large recent Homo sapiens innominates. This observation increases the likelihood that some of the additional specimens from Broken Hill are indeed of comparable antiquity to the hominid cranium and extends the range of hominids in which the feature has been recognised.     

Species-specific albumin in fossil bones from Orce, Granada, Spain

A skull fragment (VM-0) from Orce, Granada, Spain, dated palaeomagnetically at about 1.6 Myr, is thought by some palaeontologist to be hominid, while others maintain it is equid. If hominid, it would be by far the oldest evidence ofHomo in Europe. Immunological studies on residual albumin in this fossil were carried out independently, and with different immunological methods, at the University of California, San Francisco (radioimmunoassay), and at the University of Granada, Spain (enzyme immunoassay). Other fossils attributed to hominids also studied wereVM1960 from Venta Micena, andCV-1 from Cueva Victoria, Murcia, Spain. Undisputed equid and bovid fossils from the same deposits and dated to a similar period as the Orce skull were also analyzed. Our results showed that species-specific albumin can be detected in 1.6 Myr-old hominid, equid and bovid fossils. The albumin from the Orce skull fragment and fromVM-1960 was immunologically closer to human albumin. These findings support the contention that theVM-0 andVM-1960 are hominid and that members of the genusHomo occupied southern Spain 1.6 Myr ago.     

Variation in enamel development of South African fossil hominids

Dental tissues provide important insights into aspects of hominid palaeobiology that are otherwise difficult to obtain from studies of the bony skeleton. Tooth enamel is formed by ameloblasts, which demonstrate daily secretory rhythms developing tissue-specific structures known as cross striations, and longer period markings called striae of Retzius. These enamel features were studied in the molars of two well known South African hominid species, Australopithecus africanus and Paranthropus robustus. Using newly developed portable confocal microscopy, we have obtained cross striation periodicities (number of cross striations between adjacent striae) for the largest sample of hominid teeth reported to date. These data indicate a mean periodicity of seven days in these small-bodied hominids. Important differences were observed in the inferred mechanisms of enamel development between these taxa. Ameloblasts maintain high rates of differentiation throughout cervical enamel development in P. robustus but not in A. africanus. In our sample, there were fewer lateral striae of Retzius in P. robustus than in A. africanus. In a molar of P. robustus, lateral enamel formed in a much shorter time than cuspal enamel, and the opposite was observed in two molars of A. africanus. In spite of the greater occlusal area and enamel thickness of the molars of both fossil species compared with modern humans, the total crown formation time of these three fossil molars was shorter than the corresponding tooth type in modern humans. Our results provide support for previous conclusions that molar crown formation time was short in Plio-Pleistocene hominids, and strongly suggest the presence of different mechanisms of amelogenesis, and thus tooth development, in these taxa.     

A note on the KNM-ER 999 hominid femur

The KNM-ER 999 hominid femur, in light of improved knowledge of later Pleistocene human postcranial morphology and ongoing reassessments of the emergence of modern humans in Africa, appears morphologically aligned with early modern humans rather than with late archaic humans. This is reflected especially in its having a clear pilaster, a proximally positioned minimum diaphyseal breadth (thereby lacking a proximo-medial diaphyseal buttress) and an exceptionally high neck-shaft angle. In these features, and especially the fast, it shows affinities to the Levantine Qafzeh-Skhul early modern human sample. However, the uncertainty regarding its stratigraphic provenience (early Late Pleistocene versus early Holocene) makes its relevance to the emergence of modern humans in Africa tenuous.     

New Late-Pleistocene uranium–thorium and ESR dates for the Singa hominid (Sudan)

The Singa (Sudan) calvaria has been interpreted previously as a terminal Pleistocene modern human fossil, perhaps related to the Bushman of Southern Africa. Here we report new mass-spectrometric U–Th dates for the calcrete deposit enclosing the fossil teeth and the calvaria itself and new electron spin resonance (ESR) dates for associated dental materials. The new data constrain the age of the hominid to at least 133±2 ka. Together with the preferred linear uptake (LU) ESR dates, the U–Th data confirm that the intriguing mixture of modern and archaic characteristics in the Singa specimen date from isotope stage 6. Far from being a modern human fossil, it represents a rare example of an archaic African population which may have been ancestral to all modernHomo sapiens.     

Relative dental development of Upper Pleistocene hominids compared to human population variation

The relative development of permanent teeth in samples of Neandertal/archaic Homo and Early Modern/Upper Paleolithic hominids is compared to the range of variability found in three recent human samples. Both fossil hominid samples are advanced in relative M2 and M3 development compared to white French-Canadians, but only the Neandertal/archaic Homo M3 sample is advanced when compared to black southern Africans. Both fossil hominid samples are delayed in relative I1 and P3 development compared to the recent human samples. Two hypotheses concerning the significance of the advanced M3 and M2 development found in both hominid groups and southern Africans compared to French-Canadians are discussed. The first postulates that the differences in relative molar development are due simply to variation in tooth/jaw size relationships. The second postulates that the relatively advanced M3 and M2 development found in the fossil hominids and southern Africans is a correlate of their potential for advanced skeletal maturation compared to French-Canadians and other European-derived populations. It appears that dental development patterns have continued to evolve from the Upper Pleistocene to present times, and that Neandertals and Early Moderns shared similar patterns of relative dental development. © 1996 Wiley-Liss, Inc.     

Pattern profile analysis of hominid and chimpanzee hand bones

In a study designed to complement morphological research on hominid hand bones, length and width measurements of the thumb, index, and middle rays were obtained from radiographs of modern human hands. These rays are primary in precision-gripping postures and are therefore the ones most relevant for investigating evolutionary changes in fine manipulation. Pattern profile analysis allows individuals or samples to be plotted against a reference sample in standard deviation units, or Z-scores. It provides an indication of how different measurements are from modern human averages, while taking into consideration the degree of variation present within modern human samples. A pattern profile for chimpanzees is clearly distinct from humans but quite similar to that of a bonobo, demonstrating the promise of pattern analysis. Partial pattern profiles of several of the more complete early hominid bones from Hadar, Swartkrans, and Olduvai (O.H. 7) are presented and compared. Hadar bones are long and wide at midshaft relative to articular widths; both body-size effects and functional differences are likely. Thumb distal phalanges from Swartkrans and Olduvai both have relatively small base widths, but they differ in other proportions. Two first metacarpals from Swartkrans show distinct patterns. The profiles of La Ferrassie I and Shanidar IV show the characteristically large Neanderthal distal phalanges. Profiles of Skhūl IV and P?edmost III are alike in some regions with reference to modern North American white males, though they are less similar overall than are those of the two Neanderthals. © 1995 Wiley-Liss, Inc.     

How big were early hominids?

The recent discovery of new postcranial fossils, particularly associated body parts, of several Plio-Pleistocene hominids provides a new opportunity to assess body size in human evolution.¹ Body size plays a central role in the biology of animals because of its relationship to brain size, feeding behavior, habitat preference, social behavior, and much more. Unfortunately, the prediction of body weight from fossils is inherently inaccurate because skeletal size does not reflect body size exactly and because the fossils are from species having body proportions for which there are no analogues among modern species. The approach here is to find the relationship between body size and skeletal size in ape and human specimens of known body weight at death and to apply this knowledge to the hominid fossils, using a variety of statistical methods, knowledge of the associated partial skeletons of the of early hominids, formulae derived from a modern human sample, and, finally, common sense. The following modal weights for males and females emerge: Australopithecus afarensis, 45 and 29 kg; A. africanus, 41 and 30 kg; A. robustus, 40 and 32 kg; A. boisei, 49 and 34 kg; H. habilis, 52 and 32 kg. The best known African early H. erectus were much larger with weights ranging from 55 kg on up. These estimates imply that (1) in the earliest hominid species and the “robust” australopithecines body sizes remained small relative to modern standards, but between 2.0 and 1.7 m.y.a. there was a rapid increase to essentially modern body size with the appearance of Homo erectus; (2) the earliest species had a degree of body size sexual dimorphism well above that seen in modern humans but below that seen in modern gorillas and orangs which implies (along with other evidence) a social organization characterized by kin-related, multi-male groups with females who were not kin-related; (3) relative brain sizes increased through time; (4) there were two divergent trends in relative cheek-tooth size—a steady increase through time from A. afarensis to A. africanus to the “robust” australopithecines, and a decrease beginning with H. habilis to H. erectus to H. sapiens.     

Early ontogeny of the human femoral bicondylar angle

The presence of a femoral bicondylar angle consistently and significantly greater than 0° has been a hallmark of hominid bipedality, but its pattern of development has not been documented. We have therefore compiled cross-sectional data on the development of the articular bicondylar angle for a clinical sample of modern humans and of the metaphyseal bicondylar angle for two Recent human skeletal samples, one predominantly European in origin and the other Amerindian. All three samples exhibit a pattern of a bicondylar angle of 0° at birth and then a steady average increase in the angle from late in the first year postnatal, through infancy, and into the juvenile years. The two skeletal samples reach low adult values by approximately 4 years postnatal, whereas the clinical sample with a lowered activity level appears to attain consistent adult values slightly later (approximately 6 years postnatal). In addition, two modern human individuals, one nonambulatory and the other minimally ambulatory, show no and little development, respectively, of a bicondylar angle. These data, in conjunction with clinical and experimental observations on the potential and form of angular changes during epiphyseal growth, establish a high degree of potential for plasticity in the development of the human bicondylar angle and the direct association of a bipedal locomotion and (especially) posture with the developmental emergence of a human femoral bicondylar angle. © 1994 Wiley-Liss, Inc.     

First direct dating of a presumed Pleistocene hominid from China: AMS radiocarbon age of a femur from the Ordos Plateau

Human remains from the Xarusgol Valley, Ordos Plateau, northwestern China, have been considered to date to the Late Pleistocene. In order to ascertain their true age, direct AMS (14)C dating of a femur collected in the early 1920s was conducted. The results demonstrate that the femur is very young, with one sample of 'post-bomb' age and the other sample c. 200 years old. This first direct dating of a Chinese fossil hominid underscores the need to apply the same methodology to other Chinese modern human fossils currently believed to be of Pleistocene age.     

Allometric patterns of cranial bone thickness in fossil hominids

The interspecific allometry of five measures of total cranial bone thickness is examined in 10 extant catarrhine genera and two fossil hominid samples representing A. africanus and Asian H. erectus. Analysis of the modern sample shows that most interspecific variation in vault thickness can be accounted for by variation in body size. Correlation values are moderate to high (r = 0.75–0.98), and all variables exhibit positive allometry. The bone thickness:body mass relationship of modern humans broadly conforms with that of other primates. However, in the distribution of relative thickness throughout the skull, H. sapiens is distinguished by relative thickening of the parietal and extreme relative thinning of the temporal squama. The bone thickness:body mass relationship in the two early hominid species is examined using published mean body weight estimates generated from post-cranial predictor variables. A. africanus exhibits great similarity to modern humans in its relation to the catarrhine regression data and in the distribution of relative thickness throughout the skull. H. erectus also shows a modern human-like pattern in the distribution of its relative thickness; however, its bone thickness:body mass relationship is dissimilar to that displayed by all other taxa, including the other hominid species. On the basis of these results, it is suggested that the published body weight estimate assigned to H. erectus greatly underestimates actual mean body size for Asian members of this species. © 1996 Wiley-Liss, Inc.     

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.