Hominin Species Research Articles

The role of culture in early expansions of humans – A new research center

During the last two million years, the geographic range of hominin taxa dispersed in several waves from its original African homeland to encompass Asia and Europe – and possibly back. Of these hominin species, only anatomically and behaviorally modern man, Homo sapiens sapiens, has been able to overcome the impediments imposed by the physical geography and environmental conditions of this planet. Within a few tens of thousands of years, modern man successfully inhabited the globe, settling in Australia, the Americas and even the polar regions. Organisms are bound by specific ranges of environmental conditions to which they have adapted over many successive generations. As the spatial distribution of these environmental conditions changes over time, well adapted populations can follow an ameliorating climate up to its boundaries. However, relative deterioration of the basic conditions can lead to a narrowing of range, whether based on climate change, availability of food, predator abundance, or perhaps even the appearance of epidemics, parasites and diseases. Expansions beyond such boundaries are possible through either biological adaptation to the changing environmental conditions or through innovative cultural adaptation. In this regard, the knowledge of an organism’s specific habitat, distinguished through climatic factors, diverse physical geographical and ecological attributes, as well as the spatial and temporal development of that habitat, constitutes the minimum requirement necessary to understand a species’ expansion into new territory. The scientific scope of the new research center, ‘‘The Role of Culture in Early Expansions of Humans’’ (Fig. 1), which is funded by the Heidelberg Academy of Sciences and Humanities and projected for a duration of 20 years, seeks to reconstruct the spatial and temporal patterns of the expansion of hominins between three million and 20,000 years ago in Africa and Eurasia (e.g. Schrenk 2008; Serangeli and Bolus, 2008). Other facets of the research center explore potential expansion routes, biological mechanisms and the cultural potential with which our ancestors sustained themselves in new environments. The main goal is to find reasons for different hominin expansions. Implicit in the current working hypothesis is the assumption that the impact of changing environmental conditions on hominin dispersal decreased as the importance of cultural and technological innovations grew. The creation of a web-based database will lay the foundation for the scientific investigation into the earlier expansions of humankind. The database will integrate vegetational history, paleontology, climatic record and geophysical information in standardized and

Liang Bua Homo floresiensis mandibles and mandibular teeth: a contribution to the comparative morphology of a new hominin species

In 2004, a new hominin species, Homo floresiensis, was described from Late Pleistocene cave deposits at Liang Bua, Flores. H. floresiensis was remarkable for its small body-size, endocranial volume in the chimpanzee range, limb proportions and skeletal robusticity similar to Pliocene Australopithecus, and a skeletal morphology with a distinctive combination of symplesiomorphic, derived, and unique traits. Critics of H. floresiensis as a novel species have argued that the Pleistocene skeletons from Liang Bua either fall within the range of living Australomelanesians, exhibit the attributes of growth disorders found in modern humans, or a combination of both. Here we describe the morphology of the LB1, LB2, and LB6 mandibles and mandibular teeth from Liang Bua. Morphological and metrical comparisons of the mandibles demonstrate that they share a distinctive suite of traits that place them outside both the H. sapiens and H. erectus ranges of variation. While having the derived molar size of later Homo, the symphyseal, corpus, ramus, and premolar morphologies share similarities with both Australopithecus and early Homo. When the mandibles are considered with the existing evidence for cranial and postcranial anatomy, limb proportions, and the functional anatomy of the wrist and shoulder, they are in many respects closer to African early Homo or Australopithecus than to later Homo. Taken together, this evidence suggests that the ancestors of H. floresiensis left Africa before the evolution of H. erectus, as defined by the Dmanisi and East African evidence.

Homo floresiensis from head to toe

Fossils of tiny ancient humans, found on the island of Flores, have provoked much debate and speculation. Evidence that they are a real species comes from analyses of the foot and also — more surprisingly — of dwarf hippos. The diminutive hominin Homo floresiensis, first described in Nature in 2004, lived on the island of Flores in Indonesia until about 14,000 years ago. The cover shows the partial skeleton of the type specimen, LB1. It preserves enough material to permit partial assembly of the foot, and that assembly is now reported by Jungers et al. LB1's foot had human-like fully adducted big toes, but they were longer relative to the rest of the lower limb than in modern humans, instead resembling some apes. The idea that H. floresiensis was a diminutive hominin met with some scepticism, in particular its small brain size was attributed by some to pathology. New work by Eleanor Weston and Adrian Lister shows that the brains of extinct dwarf hippos from Madagascar were disproportionately a lot smaller than those of their mainland relatives. This supports suggestions that island dwarfism breaks the 'rule' that body size reduction in mammals is accompanied by only a moderate reduction in brain size. In News & Views, Daniel Lieberman discusses these papers and a special issue of The Journal of Human Evolution, and concludes that H. floresiensis probably is a bone fide — and very interesting — species of hominin.

The foot of Homo floresiensis

Homo floresiensis is an endemic hominin species that occupied Liang Bua, a limestone cave on Flores in eastern Indonesia, during the Late Pleistocene epoch. The skeleton of the type specimen (LB1) of H. floresiensis includes a relatively complete left foot and parts of the right foot. These feet provide insights into the evolution of bipedalism and, together with the rest of the skeleton, have implications for hominin dispersal events into Asia. Here we show that LB1's foot is exceptionally long relative to the femur and tibia, proportions never before documented in hominins but seen in some African apes. Although the metatarsal robusticity sequence is human-like and the hallux is fully adducted, other intrinsic proportions and pedal features are more ape-like. The postcranial anatomy of H. floresiensis is that of a biped, but the unique lower-limb proportions and surprising combination of derived and primitive pedal morphologies suggest kinematic and biomechanical differences from modern human gait. Therefore, LB1 offers the most complete glimpse of a bipedal hominin foot that lacks the full suite of derived features characteristic of modern humans and whose mosaic design may be primitive for the genus Homo. These new findings raise the possibility that the ancestor of H. floresiensis was not Homo erectus but instead some other, more primitive, hominin whose dispersal into southeast Asia is still undocumented.

The prehistory of compassion

Beyond the biological traits that differentiate ancient species of hominins from extant humans, differences in social organization remain an important yet difficult issue to assess. Psychological features and interindividual relations, in particular, are among the least accessible aspects of past behaviors. Although the fossil and archaeological records do not easily allow us to tackle these kinds of issues, contrasting views on the cognitive capabilities and behavioral sophistication of fossil human species have often been expressed. In this issue of PNAS, Gracia et al. (1) provide new evidence on the survival of an abnormal individual with possible cognitive deficits from a group of pre-Neandertal Pleistocene hunter-gatherers, currently assigned to a geological age of >500 ka. The cranium SH14 from the Sima de los Huesos (Sierra de Atapuerca, Spain) is the earliest documented case of human neurocranial and brain deformity in the fossil record to date. Despite her/his pathological condition, this individual was not rejected at birth and survived until at least 5 years of age, apparently receiving the same attention as other children from the group.

LB1’s virtual endocast, microcephaly, and hominin brain evolution

Earlier observations of the virtual endocast of LB1, the type specimen for Homo floresiensis, are reviewed, extended, and interpreted. Seven derived features of LB1's cerebral cortex are detailed: a caudally-positioned occipital lobe, lack of a rostrally-located lunate sulcus, a caudally-expanded temporal lobe, advanced morphology of the lateral prefrontal cortex, shape of the rostral prefrontal cortex, enlarged gyri in the frontopolar region, and an expanded orbitofrontal cortex. These features indicate that LB1's brain was globally reorganized despite its ape-sized cranial capacity (417 cm 3). Neurological reorganization may thus form the basis for the cognitive abilities attributed to H. floresiensis. Because of its tiny cranial capacity, some workers think that LB1 represents a Homo sapiens individual that was afflicted with microcephaly, or some other pathology, rather than a new species of hominin. We respond to concerns about our earlier study of microcephalics compared with normal individuals, and reaffirm that LB1 did not suffer from this pathology. The intense controversy about LB1 reflects an older continuing dispute about the relative evolutionary importance of brain size versus neurological reorganization. LB1 may help resolve this debate and illuminate constraints that governed hominin brain evolution.

Can the Archaeology of Manual Specialization Tell Us Anything About Language Evolution? A Survey of the State of Play

In this review and position paper we explore the neural substrates for manual specialization and their possible connection with language and speech. We focus on two contrasting hypotheses of the origins of language and manual specialization: the language-first scenario and the tool-use-first scenario. Each one makes specific predictions about hand-use in non-human primates, as well as about the necessity of an association between speech adaptations and population-level right-handedness in the archaeological and fossil records. The concept of handedness is reformulated for archaeologists in terms of manual role specialization, using Guiard's model of asymmetric bimanual coordination. This focuses our attention on skilled bimanual tasks in which both upper limbs play complementary roles. We review work eliciting non-human primate hand preferences in co-ordinated bimanual tasks, and relevant archaeological data for estimating the presence or absence of a population-level bias to the right hand as the manipulator in extinct hominin species and in the early prehistory of our own species.

“Lucy” redux: A review of research on Australopithecus afarensis

In the 1970s, mid-Pliocene hominin fossils were found at the sites of Hadar in Ethiopia and Laetoli in Tanzania. These samples constituted the first substantial evidence for hominins older than 3.0 Ma and were notable for some remarkable discoveries, such as the "Lucy" partial skeleton and the abundant remains from the A.L. 333 locality at Hadar and the hominin footprint trail at Laetoli. The Hadar and Laetoli fossils were ultimately assigned to the novel hominin species Australopithecus afarensis, which at the time was the most plesiomorphic and geologically ancient hominin taxon. The discovery and naming of A. afarensis coincided with important developments in theory and methodology in paleoanthropology; in addition, important fossil and genetic discoveries were changing expectations about hominin divergence dates from extant African apes. This coincidence of events ensured that A. afarensis figured prominently in the last 30 years of paleoanthropological research. Here, the 30+ year history of discovery, analysis, and interpretation of A. afarensis and its contexts are summarized and synthesized. Research on A. afarensis continues and subject areas in which further investigation is needed to resolve ongoing debates regarding the paleobiology of this species are highlighted.

Size, shape, and asymmetry in fossil hominins: the status of the LB1 cranium based on 3D morphometric analyses

The unique set of morphological characteristics of the Liang Bua hominins ( Homo floresiensis) has been attributed to explanations as diverse as insular dwarfism and pathological microcephaly. This study examined the relationship between cranial size and shape across a range of hominin and African ape species to test whether or not cranial morphology of LB1 is consistent with the basic pattern of static allometry present in these various taxa. Correlations between size and 3D cranial shape were explored using principal components analysis in shape space and in Procrustes form space. Additionally, patterns of static allometry within both modern humans and Plio-Pleistocene hominins were used to simulate the expected cranial shapes of each group at the size of LB1. These hypothetical specimens were compared to LB1 both visually and statistically. Results of most analyses indicated that LB1 best fits predictions for a small specimen of fossil Homo but not for a small modern human. This was especially true for analyses of neurocranial landmarks. Results from the whole cranium were less clear about the specific affinities of LB1, but, importantly, demonstrated that aspects of facial morphology associated with smaller size converge on modern human morphology. This suggests that facial similarities between LB1 and anatomically modern humans may not be indicative of a close relationship. Landmark data collected from this study were also used to test the degree of cranial asymmetry in LB1. These comparisons indicated that the cranium is fairly asymmetrical, but within the range of asymmetry exhibited by modern humans and all extant African ape species. Compared to other fossil specimens, the degree of asymmetry in LB1 is moderate and readily explained by the taphonomic processes to which all fossils are subject. Taken together, these findings suggest that H. floresiensis was most likely the diminutive descendant of a species of archaic Homo, although the details of this evolutionary history remain obscure.

‘Whatever turns you on’: a response to Anna Machin, ‘Why handaxes just aren't that sexy’

Evaluating theories and testing hypotheses that relate to the no-longer observable behaviour of hominin species which have no close analogue in the modern world is an obvious challenge. Machin argues that Kohn & Mithen (1999) did not do so in a sufficiently rigorous manner concerning their so-called 'Sexy Handaxe Theory' (SHT). She is right, of course. Indeed how could it be otherwise when there is always room for improvement by subjecting previously published ideas to newly available data-sets and new types of critical thinking. Machin provides some valuable arguments, bringing together a probably unparalleled breadth of knowledge about the archaeological record, evolutionary theory and sexual selection. Ultimately, however, I am not persuaded that she makes handaxes any less sexy than they had previously appeared.

Hominin life history: reconstruction and evolution.

In this review we attempt to reconstruct the evolutionary history of hominin life history from extant and fossil evidence. We utilize demographic life history theory and distinguish life history variables, traits such as weaning, age at sexual maturity, and life span, from life history-related variables such as body mass, brain growth, and dental development. The latter are either linked with, or can be used to make inferences about, life history, thus providing an opportunity for estimating life history parameters in fossil taxa. We compare the life history variables of modern great apes and identify traits that are likely to be shared by the last common ancestor of Pan-Homo and those likely to be derived in hominins. All great apes exhibit slow life histories and we infer this to be true of the last common ancestor of Pan-Homo and the stem hominin. Modern human life histories are even slower, exhibiting distinctively long post-menopausal life spans and later ages at maturity, pointing to a reduction in adult mortality since the Pan-Homo split. We suggest that lower adult mortality, distinctively short interbirth intervals, and early weaning characteristic of modern humans are derived features resulting from cooperative breeding. We evaluate the fidelity of three life history-related variables, body mass, brain growth and dental development, with the life history parameters of living great apes. We found that body mass is the best predictor of great ape life history events. Brain growth trajectories and dental development and eruption are weakly related proxies and inferences from them should be made with caution. We evaluate the evidence of life history-related variables available for extinct species and find that prior to the transitional hominins there is no evidence of any hominin taxon possessing a body size, brain size or aspects of dental development much different from what we assume to be the primitive life history pattern for the Pan-Homo clade. Data for life history-related variables among the transitional hominin grade are consistent and none agrees with a modern human pattern. Aside from mean body mass, adult brain size, crown and root formation times, and the timing and sequence of dental eruption of Homo erectus are inconsistent with that of modern humans. Homo antecessor fossil material suggests a brain size similar to that of Homo erectus s. s., and crown formation times that are not yet modern, though there is some evidence of modern human-like timing of tooth formation and eruption. The body sizes, brain sizes, and dental development of Homo heidelbergensis and Homo neanderthalensis are consistent with a modern human life history but samples are too small to be certain that they have life histories within the modern human range. As more life history-related variable information for hominin species accumulates we are discovering that they can also have distinctive life histories that do not conform to any living model. At least one extinct hominin subclade, Paranthropus, has a pattern of dental life history-related variables that most likely set it apart from the life histories of both modern humans and chimpanzees.

Cranial base evolution within the hominin clade

The base of the cranium (i.e. the basioccipital, the sphenoid and the temporal bones) is of particular interest because it undergoes significant morphological change within the hominin clade, and because basicranial morphology features in several hominin species diagnoses. We use a parsimony analysis of published cranial and dental data to predict the cranial base morphology expected in the hypothetical last common ancestor of the Pan-Homo clade. We also predict the primitive condition of the cranial base for the hominin clade, and document the evolution of the cranial base within the major subclades within the hominin clade. This analysis suggests that cranial base morphology has continued to evolve in the hominin clade, both before and after the emergence of the genus Homo.

Functional ecology and evolution of hominoid molar enamel thickness: Pan troglodytes schweinfurthii and Pongo pygmaeus wurmbii

The divergent molar characteristics of Pan troglodytes and Pongo pygmaeus provide an instructive paradigm for examining the adaptive form-function relationship between molar enamel thickness and food hardness. Although both species exhibit a categorical preference for ripe fruit over other food objects, the thick enamel and crenulated occlusal surface of Pongo molar teeth predict a diet that is more resistant to deformation (hard) and fracture (tough) than the diet of Pan. We confirm these predictions with behavioral observations of Pan troglodytes schweinfurthii and Pongo pygmaeus wurmbii in the wild and describe the mechanical properties of foods utilized during periods when preferred foods are scarce. Such fallback foods may have exerted a selective pressure on tooth evolution, particularly molar enamel thinness, which is interpreted as a functional adaptation to seasonal folivory and a derived character trait within the hominoid clade. The thick enamel and crenulated occlusal surface of Pongo molars is interpreted as a functional adaptation to the routine consumption of relatively tough and hard foods. We discuss the implications of these interpretations for inferring the diet of hominin species, which possessed varying degrees of thick molar enamel. These data, which are among the first reported for hominoid primates, fill an important empirical void for evaluating the mechanical plausibility of putative hominin food objects.

Paleoecological patterns at the Hadar hominin site, Afar Regional State, Ethiopia

Reconstructing paleoecological patterns associated with hominin taxa, such as Australopithecus afarensis, is important for understanding possible evolutionary mechanisms involved in extinction and speciation events. It is critical to identify local, regional, or pan-African causal factors because patterns at these different levels may affect separate populations of the same species of hominin in unique ways. Habitat reconstructions of 12 submembers of the Hadar and Busidima formations (∼ 3.8–2.35 Ma) are presented here along with faunal differences in these submembers through time. Habitats with medium density tree and bush cover dominated the landscape through much of the earlier time period in the Hadar Formation. The lowermost Sidi Hakoma Member is the most closed habitat. The Denen Dora Member shows the influence of frequent floodplain edaphic grasslands with high abundances of reducin bovids. There is an influx of ungulates in the Kada Hadar Member (∼3.2–∼2.96 Ma) that indicates a more arid habitat populated by mammals that were recovered from earlier deposits further south in Ethiopia and Kenya. In the younger deposits from the Busidima Formation at Hadar, the landscape was open wooded grassland with some floodplain environments. The fossil assemblages from the Busidima Formation show a substantial species turnover. Although high numbers of A. afarensis specimens are associated with the lower Sidi Hakoma Member, they clearly inhabited a variety of habitats throughout the entire Hadar Formation. Australopithecus afarensis from Laetoli through Hadar times appears to have been a eurytopic species.

Extended Male Growth in a Fossil Hominin Species

In primates that are highly sexually dimorphic, males often reach maturity later than females, and young adult males do not show the size, morphology, and coloration of mature males. Here we describe extended male development in a hominin species, Paranthropus robustus. Ranking a large sample of facial remains on the basis of dental wear stages reveals a difference in size and robusticity between young adult and old adult males. Combined with estimates of sexual dimorphism, this pattern suggests that male reproductive strategy focused on monopolizing groups of females, in a manner similar to that of silverback gorillas. However, males appear to have borne a substantial cost in the form of high rates of predation.

Tectonics, orbital forcing, global climate change, and human evolution in Africa: introduction to the African paleoclimate special volume

The late Cenozoic climate of Africa is a critical component for understanding human evolution. African climate is controlled by major tectonic changes, global climate transitions, and local variations in orbital forcing. We introduce the special African Paleoclimate Issue of the Journal of Human Evolution by providing a background for and synthesis of the latest work relating to the environmental context for human evolution. Records presented in this special issue suggest that the regional tectonics, appearance of C 4 plants in East Africa, and late Cenozoic global cooling combined to produce a long-term drying trend in East Africa. Of particular importance is the uplift associated with the East African Rift Valley formation, which altered wind flow patterns from a more zonal to more meridinal direction. Results in this volume suggest a marked difference in the climate history of southern and eastern Africa, though both are clearly influenced by the major global climate thresholds crossed in the last 3 million years. Papers in this volume present lake, speleothem, and marine paleoclimate records showing that the East African long-term drying trend is punctuated by episodes of short, alternating periods of extreme wetness and aridity. These periods of extreme climate variability are characterized by the precession-forced appearance and disappearance of large, deep lakes in the East African Rift Valley and paralleled by low and high wind-driven dust loads reaching the adjacent ocean basins. Dating of these records show that over the last 3 million years such periods only occur at the times of major global climatic transitions, such as the intensification of Northern Hemisphere Glaciation (2.7–2.5 Ma), intensification of the Walker Circulation (1.9–1.7 Ma), and the Mid-Pleistocene Revolution (1–0.7 Ma). Authors in this volume suggest this onset occurs as high latitude forcing in both Hemispheres compresses the Intertropical Convergence Zone so that East Africa becomes locally sensitive to precessional forcing, resulting in rapid shifts from wet to dry conditions. These periods of extreme climate variability may have provided a catalyst for evolutionary change and driven key speciation and dispersal events amongst mammals and hominins in Africa. In particular, hominin species seem to differentially originate and go extinct during periods of extreme climate variability. Results presented in this volume may represent the basis of a new theory of early human evolution in Africa.

Implications of new early Homo fossils from Ileret, east of Lake Turkana, Kenya

Sites in eastern Africa have shed light on the emergence and early evolution of the genus Homo. The best known early hominin species, H. habilis and H. erectus, have often been interpreted as time-successive segments of a single anagenetic evolutionary lineage. The case for this was strengthened by the discovery of small early Pleistocene hominin crania from Dmanisi in Georgia that apparently provide evidence of morphological continuity between the two taxa. Here we describe two new cranial fossils from the Koobi Fora Formation, east of Lake Turkana in Kenya, that have bearing on the relationship between species of early Homo. A partial maxilla assigned to H. habilis reliably demonstrates that this species survived until later than previously recognized, making an anagenetic relationship with H. erectus unlikely. The discovery of a particularly small calvaria of H. erectus indicates that this taxon overlapped in size with H. habilis, and may have shown marked sexual dimorphism. The new fossils confirm the distinctiveness of H. habilis and H. erectus, independently of overall cranial size, and suggest that these two early taxa were living broadly sympatrically in the same lake basin for almost half a million years.

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.

The use of cranial variables for the estimation of body mass in fossil hominins

Estimating body mass/size/weight remains a crucial precursor to the evaluation of relative brain size and to achieving an understanding of brain evolution in fossil species. Despite the obvious close association between the metrics of postcranial elements and body mass a number of factors combine to reduce their utility. This study examines the feasibility of cranial variables for predicting body mass. The use of traditional regression procedures, independent contrasts analysis, and variance partitioning all support the hypothesis that cranial variables are correlated with body mass even when taking phylogeny into account, with r values typically ranging between 0.52 and 0.98. Body mass estimates derived for fossil hominins using cranial variables are similar to those obtained from previous studies using either cranial or postcranial elements. In particular, upper facial breadth and orbital height display strong predictive capability. Average body masses derived from Least Squares Regression (LSR) equations were used to calculate estimates of body mass for three hominin species. This resulted in estimates of between 30 kg and 47 kg for Australopithecus africanus, 48 kg and 52 kg for Paranthropus robustus, and 75 kg for Homo neanderthalensis. It is proposed that regression equations derived for the order primates are used to estimate body mass for archaic hominins, while hominoid based equations are most suited for Homo.

Variation and diversity in Homo erectus: a 3D geometric morphometric analysis of the temporal bone

Although the level of taxonomic diversity within the fossil hominin species Homo erectus ( sensu lato) is continually debated, there have been relatively few studies aiming to quantify the morphology of this species. Instead, most researchers have relied on qualitative descriptions or the evaluation of nonmetric characters, which in many cases display continuous variation. Also, only a few studies have used quantitative data to formally test hypotheses regarding the taxonomic composition of the “ erectus” hypodigm. Despite these previous analyses, however, and perhaps in part due to these varied approaches for assessing variation within specimens typically referred to H. erectus ( sensu lato) and the general lack of rigorous statistical testing of how variation within this taxon is partitioned, there is currently little consensus regarding whether this group is a single species, or whether it should instead be split into separate temporal or geographically delimited taxa. In order to evaluate possible explanations for variation within H. erectus, we tested the general hypothesis that variation within the temporal bone morphology of H. erectus is consistent with that of a single species, using great apes and humans as comparative taxa. Eighteen three-dimensional (3D) landmarks of the temporal bone were digitized on a total of 520 extant and fossil hominid crania. Landmarks were registered by Generalized Procrustes Analysis, and Procrustes distances were calculated for comparisons of individuals within and between the extant taxa. Distances between fossil specimens and between a priori groupings of fossils were then compared to the distances calculated within the extant taxa to assess the variation within the H. erectus sample relative to that of known species, subspecies, and populations. Results of these analyses indicate that shape variation within the entire H. erectus sample is generally higher than extant hominid intraspecific variation, and putative H. ergaster specimens are significantly different from other specimens in H. erectus ( sensu lato). However, shape distances within geographical groups of H. erectus are also high, and OH 9 and Dmanisi 2280 are morphologically distinct from the Koobi Fora specimens that are sometimes classified as H. ergaster. These findings suggest that, although H. erectus may be composed of multiple species, the differentiation is complex, and specimens cannot easily be grouped geographically or chronologically. Consequently, more complicated scenarios seeking to explain the observed variation within H. erectus must be considered.