Human Fossil Record Research Articles

“Small Size” in the Philippine Human Fossil Record: Is It Meaningful for a Better Understanding of the Evolutionary History of the Negritos?

“Pygmy populations” are recognized in several places over the world, especially in Western Africa and in Southeast Asia (Philippine “negritos,” for instance). Broadly defined as “small-bodied Homo sapiens” (compared with neighboring populations), their origins and the nature of the processes involved in the maintenance of their phenotype over time are highly debated. Major results have been recently obtained from population genetics on present-day negrito populations, but their evolutionary history remains largely unresolved. We present and discuss the Upper Pleistocene human remains recovered from Tabon Cave and Callao Cave in the Philippines, which are potentially highly relevant to these research questions. Human fossils have been recovered in large numbers from Tabon Cave (Palawan Island) but mainly from reworked and mixed sediments from several archaeological layers. We review and synthesize the long and meticulous collaborative work done on the archives left from the 1960s excavations and on the field. The results demonstrate the long history of human occupations in the cave, since at least ~30,000 BP. The examination of the Tabon human remains shows a large variability: large and robust for one part of the sample, and small and gracile for the other part. The latter would fit quite comfortably within the range of variation of Philippine negritos. Farther north, on Luzon Island, the human third metatarsal recently recovered from Callao Cave and dated to ~66,000 BP is now the oldest direct evidence of human presence in the Philippines. Previous data show that, compared with H. sapiens (including Philippine negritos), this bone presents a very small size and several unusual morphological characteristics. We present a new analytical approach using three-dimensional geometric morphometrics for comparing the Callao fossil to a wide array of extant Asian mammals, including nonhuman primates and H. sapiens. The results demonstrate that the shape of the Callao metatarsal is definitely closer to humans than to any other groups. The fossil clearly belongs to the genus Homo; however, it remains at the margin of the variation range of H. sapiens. Because of its great antiquity and the presence of another diminutive species of the genus Homo in the Wallace area during this time period (H. floresiensis), we discuss here in detail the affinities and potential relatedness of the Callao fossil with negritos that are found today on Luzon Island.

“Small Size” in the Philippine Human Fossil Record: Is It Meaningful for a Better Understanding of the Evolutionary History of the Negritos?

“Small Size” in the Philippine Human Fossil Record: Is It Meaningful for a Better Understanding of the Evolutionary History of the Negritos?

Dentoalveolar paleopathology of the early modern humans from Zhirendong, South China

The fragmentary early Late Pleistocene, early modern human remains from Zhirendong, south China, present a suite of dentoalveolar pathologies and anomalies. The lesions include lower molar buccal alveolar resorption (Zhiren 1), massive dental caries in a mandibular molar associated with hypercementosis (Zhiren 2), and bilateral mesial premolar (P3) periapical lesions (granulomata with a probable left abscess) (Zhiren 3). The Zhiren 3 periapical lesions, given their bilaterality and the non-pathological incisor and canine alveoli, suggest dens evaginatus, although absence of the Zhiren 3 dentition prevents full evaluation of this diagnosis. These periodontal abnormalities join a number of similar lesions in Pleistocene humans, of varying severity. The carious lesion is noteworthy, given the rarity of them in the Pleistocene human fossil record. In addition, Zhiren 3 exhibits unilateral P3 rotation and bilateral I1 rotation (winging).

Species-specific responses of Late Quaternary megafauna to climate and humans

Despite decades of research, the roles of climate and humans in driving the dramatic extinctions of large-bodied mammals during the Late Quaternary period remain contentious. Here we use ancient DNA, species distribution models and the human fossil record to elucidate how climate and humans shaped the demographic history of woolly rhinoceros, woolly mammoth, wild horse, reindeer, bison and musk ox. We show that climate has been a major driver of population change over the past 50,000 years. However, each species responds differently to the effects of climatic shifts, habitat redistribution and human encroachment. Although climate change alone can explain the extinction of some species, such as Eurasian musk ox and woolly rhinoceros, a combination of climatic and anthropogenic effects appears to be responsible for the extinction of others, including Eurasian steppe bison and wild horse. We find no genetic signature or any distinctive range dynamics distinguishing extinct from surviving species, emphasizing the challenges associated with predicting future responses of extant mammals to climate and human-mediated habitat change.

The relative efficacy of functional and developmental cranial modules for reconstructing global human population history

This study tests the relative efficacy of human cranial modules, defined on the basis of developmental and functional criteria, for reconstructing neutral genetic population history. Specifically, two hypotheses were tested: 1) The "basicranial hypothesis" predicts that the endochondrally ossifying basicranium will be more reliable for reconstructing population history than intramembranously ossifying regions of the human cranium. This is based on the assumption that early ossification of the basicranium and its distinct functional constraints produce a cranial structure that is relatively immune to non-neutral evolutionary forces. 2) The "single function hypothesis" predicts that cranial regions associated with a single (sensory) function are less reliable indicators of neutral genetic history. Here the prediction is based on the logic that complex, multi-functional, integrated cranial regions are less likely toexhibit homoplasy and, therefore, provide a more accurate morphological proxy for genetic relationships. The congruence between craniometric affinity matrices and neutral genetic population matrices based on autosomal microsatellite and classical markers was assessed using a series of Mantel and Dow-Cheverud tests. The results did not support the predictions of the "basicranial hypothesis," as the endochondrally ossifying basicranium was not significantly more congruent with the genetic data than intramembraneously ossifying modules. Moreover, although the results provided some support for the "single function hypothesis," defining cranial modules on the basis of anatomical or functional complexity did not provide a consistent means of predicting their phylogenetic efficacy. These results have important implications for building an accurate inference model of cranial evolution in the human fossil record.

Middle Pleistocene lower back and pelvis from an aged human individual from the Sima de los Huesos site, Spain

We report a nearly complete lumbar spine from the Middle Pleistocene site of the Sima de los Huesos (SH) that is assigned to the previously published SH male Pelvis 1 [Arsuaga JL, et al. (1999). Nature 399: 255-258]. The "SH Pelvis 1 individual" is a unique nearly complete lumbo-pelvic complex from the human Middle Pleistocene fossil record, and offers a rare glimpse into the anatomy and past lifeways of Homo heidelbergensis. A revised reconstruction of Pelvis 1, together with the current fossil evidence, confirms our previous hypothesis that the morphology of this pelvis represents the primitive pattern within the genus Homo. Here we argue that this primitive pattern is also characterized by sexual dimorphism in the pelvic canal shape, implying complicated deliveries. In addition, this individual shows signs of lumbar kyphotic deformity, spondylolisthesis, and Baastrup disease. This suite of lesions would have postural consequences and was most likely painful. As a result, the individual's daily physical activities would have been restricted to some extent. Reexamination of the age-at-death agrees with this individual being over 45 y old, relying on the modern human pattern of changes of the articular surfaces of the os coxae. The presence of degenerative pathological lesions and the advanced age-at-death of this individual make it the most ancient postcranial evidence of an aged individual in the human fossil record. Additional nonpathological SH lumbo-pelvic remains are consistent with previous hypotheses, suggesting a less-pronounced sagittal spinal curvature in Neandertals compared with Homo sapiens.

Evolution of middle-late Pleistocene human cranio-facial form: A 3-D approach

The classification and phylogenetic relationships of the middle Pleistocene human fossil record remains one of the most intractable problems in paleoanthropology. Several authors have noted broad resemblances between European and African fossils from this period, suggesting a single taxon ancestral to both modern humans and Neanderthals. Others point out ‘incipient’ Neanderthal features in the morphology of the European sample and have argued for their inclusion in the Neanderthal lineage exclusively, following a model of accretionary evolution of Neanderthals. We approach these questions using geometric morphometric methods which allow the intuitive visualization and quantification of features previously described qualitatively. We apply these techniques to evaluate proposed cranio-facial ‘incipient’ facial, vault, and basicranial traits in a middle-late Pleistocene European hominin sample when compared to a sample of the same time depth from Africa. Some of the features examined followed the predictions of the accretion model and relate the middle Pleistocene European material to the later Neanderthals. However, although our analysis showed a clear separation between Neanderthals and early/recent modern humans and morphological proximity between European specimens from OIS 7 to 3, it also shows that the European hominins from the first half of the middle Pleistocene still shared most of their cranio-facial architecture with their African contemporaries.

Thirty-Five Years of Paleoanthropology

When we published our first joint paper on human evolution back in early 1975 (Eldredge and Tattersall 1975), the world of human evolution was a very different place. For a start, it was populated by a much smaller cast of characters. Not only was paleoanthropology itself a much tinier enterprise than it is today, but the sum total of hominid fossils known was hugely smaller. The riches of the Turkana Basin in northern Kenya had only just begun to be explored. The iconic fossil “Lucy” had yet to be found in Ethiopia, and indeed, the Hadar area from which she came had only just been recognized for the palaeoanthropological treasure trove it has turned out to be. Now near-mythic names such as Atapuerca in Spain and Dmanisi in Georgia were yet to become part of the paleoanthropological vocabulary. And such lesser but nonetheless significant names as Ceprano, Drimolen, and Buia had still to fall even on professional ears. Yet even more foreign to modern eyes than the small fossil pond—today we can no longer complain about an “inadequate” human fossil record, much as we’d like to have more—was the prevailing way of looking at the human evolutionary story. Back in the 1970s, the view of the hominid fossil record was fairly unrelentingly minimalist. Under the sway of the Evolutionary Synthesis as it was imported into paleoanthropology in the early 1950s, students of human evolution generally sought to minimize the diversity they perceived in the known record. A single lineage struggling like a green tendril toward the light was the preferable view, and even though by the late 1960s discoveries in Kenya had rendered the strictest version of this model untenable (the “robust” australopiths had to be recognized as a side branch), there was still a general desire to simplify the picture as much as possible. This was, of course, an example of what might be called “human exceptionalism”: Since there is undeniably only one species of human on the planet today, there is an understandable desire to reconstruct our phylogeny by projecting Homo sapiens back into the past through a chain of increasingly primitive extinct species. And although among primates (and mammals) in general there is an unmistakable signal in the fossil record as well as in the modern fauna of diversity— successful forms tend to become more diverse, not to strive toward a particular goal—hominids were regarded as an exception because they possessed culture, an elusive expression of behavioral complexity that “broadened” their ecological niche to the extent that no more than one species of our kind could exist at any one time. When we began our joint investigations of human evolution in the early 1970s, the “cladistic revolution” was in full swing at least in certain corners of the American Museum of Natural History, where we both worked. And we were both highly sensitive to zoological diversity, one of us as a student of trilobites, a highly varied group of Paleozoic invertebrates, and the other as an aficionado of the lemurs, an insular group of primates inMadagascar that has diversified to an astonishing degree. We were hardly surprised, then, to find that the hominid fossils we looked at did not conform to any of the more or less linear views of human evolution that then prevailed. In particular, the cladogram we came up with (to I. Tattersall Division of Anthropology, The American Museum of Natural History, Central Park West at 79th St., New York, NY 10024, USA

Reconstructing human evolution: Achievements, challenges, and opportunities

This contribution reviews the evidence that has resolved the branching structure of the higher primate part of the tree of life and the substantial body of fossil evidence for human evolution. It considers some of the problems faced by those who try to interpret the taxonomy and systematics of the human fossil record. How do you to tell an early human taxon from one in a closely related clade? How do you determine the number of taxa represented in the human clade? How can homoplasy be recognized and factored into attempts to recover phylogeny?

New perspectives on anthropoid origins

Adaptive shifts associated with human origins are brought to light as we examine the human fossil record and study our own genome and that of our closest ape relatives. However, the more ancient roots of many human characteristics are revealed through the study of a broader array of living anthropoids and the increasingly dense fossil record of the earliest anthropoid radiations. Genomic data and fossils of early primates in Asia and Africa clarify relationships among the major clades of primates. Progress in comparative anatomy, genomics, and molecular biology point to key changes in sensory ecology and brain organization that ultimately set the stage for the emergence of the human lineage.

The vertebral formula of the last common ancestor of African apes and humans

The modal number of lumbar vertebrae in modern humans is five. It varies between three and four in extant African apes (mean=3.5). Because both chimpanzees (Pan troglodytes) and gorillas (Gorilla gorilla) possess the same distributions of thoracic, lumbar, and sacral vertebrae, it has been assumed from parsimony that the last common ancestor (LCA) of African apes and humans possessed a similarly short lower back. This "short-backed LCA" scenario has recently been viewed favorably in an analysis of the intra- and interspecific variation in axial formulas observed among African apes and humans (Pilbeam, 2004. J Exp Zool 302B:241-267). However, the number of bonobo (Pan paniscus) specimens in that study was small (N=17). Here we reconsider vertebral type and number in the LCA in light of an expanded P. paniscus sample as well as evidence provided by the human fossil record. The precaudal (pre-coccygeal) axial column of bonobos differs from those of chimpanzees and gorillas in displaying one additional vertebra as well as significantly different combinations of sacral, lumbar, and thoracic vertebrae. These findings, along with the six-segmented lumbar column of early Australopithecus and early Homo, suggest that the LCA possessed a long axial column and long lumbar spine and that reduction in the lumbar column occurred independently in humans and in each ape clade, and continued after separation of the two species of Pan as well. Such an explanation is strongly congruent with additional details of lumbar column reduction and lower back stabilization in African apes.

Into Eurasia: A geometric morphometric re-assessment of the Upper Cave (Zhoukoudian) specimens

Since the discovery of the human remains from the Upper Cave of Zhoukoudian in the 1930s there has been speculation over their affinities. In particular, the degree to which the three adult crania exhibit recent East Asian morphology, as well as their degree of within-group variability, has long been debated. Several early researchers described a resemblance to East Asian populations, but these findings have been for the most part rejected by more recent metric and non-metric analyses. Nevertheless, the Upper Cave specimens have not been classified conclusively into any recent modern human population to which they have been compared, and classification results differ for each cranium. Here, the question of the affinities of Upper Cave 101 and 103, the two better-preserved crania, is examined from the perspective of the Late Pleistocene human fossil record using the methodology of 3-D geometric morphometrics. The degree of morphological variation between the two specimens is also evaluated within the context of recent population variability. Neurocranial and facial morphology are analyzed separately so as to maximize comparative samples. Results show a morphological resemblance of the Upper Cave material to Upper Paleolithic Europeans. It is proposed that the Upper Cave specimens retain important aspects of modern human ancestral morphology, and possibly share a recent common ancestral population with Upper Paleolithic Europeans, in accordance with the Single Origin model of modern human origins.

Independent evolution of knuckle-walking in African apes shows that humans did not evolve from a knuckle-walking ancestor.

Despite decades of debate, it remains unclear whether human bipedalism evolved from a terrestrial knuckle-walking ancestor or from a more generalized, arboreal ape ancestor. Proponents of the knuckle-walking hypothesis focused on the wrist and hand to find morphological evidence of this behavior in the human fossil record. These studies, however, have not examined variation or development of purported knuckle-walking features in apes or other primates, data that are critical to resolution of this long-standing debate. Here we present novel data on the frequency and development of putative knuckle-walking features of the wrist in apes and monkeys. We use these data to test the hypothesis that all knuckle-walking apes share similar anatomical features and that these features can be used to reliably infer locomotor behavior in our extinct ancestors. Contrary to previous expectations, features long-assumed to indicate knuckle-walking behavior are not found in all African apes, show different developmental patterns across species, and are found in nonknuckle-walking primates as well. However, variation among African ape wrist morphology can be clearly explained if we accept the likely independent evolution of 2 fundamentally different biomechanical modes of knuckle-walking: an extended wrist posture in an arboreal environment (Pan) versus a neutral, columnar hand posture in a terrestrial environment (Gorilla). The presence of purported knuckle-walking features in the hominin wrist can thus be viewed as evidence of arboreality, not terrestriality, and provide evidence that human bipedalism evolved from a more arboreal ancestor occupying the ecological niche common to all living apes.

The paleoanthropology of Greece

AbstractEuropean paleoanthropology and paleolithic archeology were already well‐established by the early twentieth century. The human fossil record from this continent is the longest known and perhaps most intensively studied. Nonetheless, important gaps remain to this day in the map of Pleistocene Europe; perhaps the most glaring of these is located in the southeastern corner of the continent. This region's record is critical for addressing questions about the course of human evolution in Europe because its geographic position lends it a dual role: on one hand, it encompasses a frequently hypothesized dispersal corridor from Africa into Europe for both archaic and early modern humans; on the other, as one of the three Mediterranean peninsulas, it acted as a refugium for plant, animal, and, most likely, human populations during glacial conditions. This article is a review of the paleoanthopological record of Greece, one of the least known in Europe.

A Celebration of Clifford Jolly’s Contribution to Biological Anthropology

Biological anthropology is a unique scientific field in which, possibly more than in other sciences, progress depends on the effective integration and synthesis of data and evidence from a wide variety of disciplines. Combining the complexity and depth of modern biology with a desire to understand the unique historical event that is the origin of our own species, significant advances in biological anthropology often develop from the creative application of new methods developed in one scientific discipline to questions posed by researchers with entirely different interests. In addition, some of the most important advances during the history of human evolutionary studies have come from individual investigators who reframed traditional questions in light of new comparative evidence drawn from the living primates. Clifford Jolly‘s long and remarkably successful career illustrates the value and impact of integrating evidence and concepts from disparate fields in the service of the broad goals of biological anthropology. He has made substantive and lasting contributions to the interpretation of the human fossil record, to both knowledge of and methods used in the study of natural populations of living primates, to laboratory procedures for investigating those primates, and certainly to the conceptual framework that biological anthropologists employ to understand evolutionary diversity within and among primate species. In order to celebrate the many significant contributions Cliff has made and continues to make to our broad field of study, and to highlight the current exciting state of biological anthropology in general, we organized a conference designed to bring together active researchers from a variety of subfields in which he has worked. This section of Current Anthropology presents a series of five papers first presented at that conference. The conference, titled “Evolutionary Anthropology at the Interface,” was a two-day event held in New York in October 2007

New data on the late Neandertals: Direct dating of the Belgian Spy fossils

In Eurasia, the period between 40,000 and 30,000 BP saw the replacement of Neandertals by anatomically modern humans (AMH) during and after the Middle to Upper Paleolithic transition. The human fossil record for this period is very poorly defined with no overlap between Neandertals and AMH on the basis of direct dates. Four new (14)C dates were obtained on the two adult Neandertals from Spy (Belgium). The results show that Neandertals survived to at least approximately 36,000 BP in Belgium and that the Spy fossils may be associated to the Lincombian-Ranisian-Jerzmanowician, a transitional techno-complex defined in northwest Europe and recognized in the Spy collections. The new data suggest that hypotheses other than Neandertal acculturation by AMH may be considered in this part of Europe.

Human uniqueness-self-interest and social cooperation

Humans are unique among all species of terrestrial history in both ecological dominance and individual properties. Many, or perhaps all, of the unique elements of this nonpareil status can be plausibly interpreted as evolutionary and strategic elements and consequences of the unprecedented intensity and scale of our social cooperation. Convincing explanation of this unique human social adaptation remains a central, unmet challenge to the scientific enterprise. We develop a hypothesis for the ancestral origin of expanded cooperative social behavior. Specifically, we present a game theoretic analysis demonstrating that a specific pattern of expanded social cooperation between conspecific individuals with conflicts of interest (including non-kin) can be strategically viable, but only in animals that possess a highly unusual capacity for conspecific violence (credible threat) having very specific properties that dramatically reduce the costs of coercive violence. The resulting reduced costs allow preemptive or compensated coercion to be an instantaneously self-interested behavior under diverse circumstances rather than in rare, idiosyncratic circumstances as in actors (animals) who do not have access to inexpensive coercive threat. Humans are apparently unique among terrestrial organisms in having evolved conspecific coercive capabilities that fulfill these stringent requirements. Thus, our results support the proposal that access to a novel capacity for projection of coercive threat might represent the essential initiating event for the evolution of a human-like pattern of social cooperation and the subsequent evolution of the diverse features of human uniqueness. Empirical evidence indicates that these constraints were, in fact, met only in our evolutionary lineage. The logic for the emergence of uniquely human cooperation suggested by our analysis apparently accounts simply for the human fossil record.

The Homo floresiensis Controversy

A completely new and unexpected quasi human species, Homo floresiensis, nicknamed the Hobbit, was described in 2004 from Liang Bua, a cave in Flores. Like many important new contributions to the human fossil record in the past, many commentators refused to believe that a new species had been discovered, and the type specimen was interpreted as a pathological modern human, usually as a microcephalic dwarf. There is no substance to these claims: close analysis shows that Homo floresiensis is not only a genuinely new species, but that its closest affinities lie with Plio-Pleistocene African species such as Homo habilis, so that it documents an earlier dispersal of hominins from Africa and had hitherto been suspected.

A geometric morphometric analysis of hominin upper first molar shape

Recent studies have revealed interesting differences in upper first molar morphology across the hominin fossil record, particularly significant between H. sapiens and H. neanderthalensis. Usually these analyses have been performed by means of classic morphometric methods, including the measurement of relative cusp areas or the angles defined between cusps. Although these studies have provided valuable information for the morphological characterization of some hominin species, we believe that the analysis of this particular tooth could be more conclusive for taxonomic assignment. In this study, we have applied geometric morphometric methods to explore the morphological variability of the upper first molar (M 1) across the human fossil record. Our emphasis focuses on the study of the phenetic relationships among the European middle Pleistocene populations (designated as H. heidelbergensis) with H. neanderthalensis and H. sapiens, but the inclusion of Australopithecus and early Homo specimens has helped us to assess the polarity of the observed traits. H. neanderthalensis presents a unique morphology characterized by a relatively distal displacement of the lingual cusps and protrusion in the external outline of a large and bulging hypocone. This morphology can be found in a less pronounced degree in the European early and middle Pleistocene populations, and reaches its maximum expression with the H. neanderthalensis lineage. In contrast, modern humans retain the primitive morphology with a square occlusal polygon associated with a round external outline.

Paleoanthropology, the Short Version

The Very Short Introduction series is just that, a series of pocket-sized (4.5× 7”) books about a wide variety of topics. It seems similar to the now-defunct French series “Que sais-je” (“what do I know,” or better “what should I know”). In this volume, Bernard Wood, a leading scholar of human evolution, presents the main theoretical background, fossil data, and interpretations of biological paleoanthropology; he specifically bypasses archaeology, which is set to appear in another “VSI” on Prehistory, but that loses one of the main integrative benefits of modern paleoanthropology. As part of the introductory/historical section, Wood presents two alternative classifications of higher primates, which will interest readers of Journal of Mammalian Evolution. In the traditional version, there are two families other than Hylobatidae (all apes in Pongidae; a subfamily Australopithecinae for all genera other than Homo within Hominidae), while there is just one in the version that includes genetic evidence; here, however, Wood is hoisted by his own petard, recognizing Ponginae, Gorillinae, and Homininae (with Panini and Hominini). The reason for recognizing an Italian sandwich as a tribe for chimps is obviously to keep our own minor clade at the currently-desired tribal level, but this leads to imbalance at the subfamily level. There is no simple answer-either humans must devolve to subtribe Hominina (homininans) or we can cheat as I have done and place Pan as Homininae indet., recognizing Gorillini and Hominini (or, of course, abandon Linnaean nomenclature altogether). About halfway through the book (chapter 5, p. 58), Wood begins to consider the human fossil record, with successive chapters on: the predecessors to Australopithecus; Australopithecus, Paranthropus, and the earliest species of Homo (which Wood here returns to that genus, despite previous ejection in Wood and Collard, 1999a,b); later species of Homo; and modern humans, including our origins and global dispersal. Each of these chapters has a “points to watch” box of current questions and controversies, as well as a site map and a typical Wood phylogram of taxon ranges without clear indications of phylogeny. Unfortunately, the size restrictions prevent the inclusion of photographs or even drawings of the fossils themselves, and morphology in general