Homo erectus has been broadly defined to include fossils from Africa, Asia, and possibly Europe, or restricted to a supposedly confined Asian clade. Recently discovered fossils of H. erectus are allowing new insights into aspects of its evolution, such as the timing and mode of the species’ emergence in Africa and its relationship to Asian populations. However, the currently available African record predating 1.0 Ma is poor, consisting of the Turkana basin, Olduvai and the more limited South African materials. Here, we describe and compare eight craniodental fossils of ~1.4 Ma recovered from Konso, Ethiopia, that we attribute to H. erectus. These include KGA10-1, one of the better-preserved H. erectus mandibular specimens known from eastern Africa, and other fragmentary dental and cranial remains. The Konso H. erectus fossils show a mosaic of primitive and derived features. These include a large and thick mandibular corpus, a moderately developed lateral prominence, a reduced premolar morphology, and a tendency for smaller relative sizes of the posterior molars compared with earlier Homo. In some dentognathic details, such as the lack of a buccolingually narrow M1 and the presence of double mental foramina, the Konso fossils differ from eastern African H. erectus of ≥1.5 Ma. The fragmentary cranial remains exhibit weak angular and occipital tori, and an apparently weak occipital flexion, as with the eastern African H. erectus examples known from ~1.65 to 1.2 Ma. The available evidence is consistent with the interpretation that African early H. erectus shows morphological continuity within the ~1.65 to 1.0 Ma time period, with relatively little morphological evolution prior to 1.4 Ma and advanced dentognathic gracility occurring sometime thereafter. The Konso evidence corroborates the hypothesis that the African H. erectus populations represent a variable but continuous evolutionary succession that was a likely source of multiple events of gene flow to the Eurasian continent.
Our view of H. erectus is vastly different today than when Pithecanthropus erectus was described in 1894. Since its synonimization into Homo, views of the species and its distribution have varied from a single, widely dispersed, polytypic species ultimately ancestral to all later Homo, to a derived, regional isolate ultimately marginal to later hominin evolution. A revised chronostratigraphic framework and recent work bearing either directly or indirectly on reconstructions of life-history patterns are reviewed here and, together with a review of the cranial and postcranial anatomy of H. erectus, are used to generate a natural history of the species. Here I argue that H. erectus is a hominin, notable for its increased body size, that originates in the latest Pliocene/earliest Pleistocene of Africa and quickly disperses into Western and Eastern Asia. It is also an increasingly derived hominin with several regional morphs sustained by intermittent isolation, particularly in Southeast Asia. This view differs from several current views, most especially that which recognizes only a single hominin species in the Pleistocene, H. sapiens, and those which would atomize H. erectus into a multiplicity of taxa. Following Jolly ([2001] Yrbk Phys Anthropol 44:177-204), the regional morphs of H. erectus may be productively viewed as geographically replacing allotaxa, rather than as the focus of unresolvable species debates. Such a view allows us to focus on the adaptations and biology of local groups, including questions of biogeographic isolation and local adaptation. A number of issues remain unresolved, including the significance of diversity in size and shape in the early African and Georgian records.
This paper investigates patterns of cranial capacity evolution in Homo erectus, early Homo sapiens, and in regional subsamples of H. erectus. Specifically, models explaining evolution of cranial capacity in these taxa are evaluated with statistical techniques developed for the analysis of time series data. Regression estimates of rates of evolution in cranial capacity are also obtained. A non-parametric test for trend suggests that cranial capacity in both H. erectus and early H. sapiens may increase significantly through time. Cranial capacity in an Asian subsample of H. erectus (comprised of Chinese and Indonesian specimens) increases significantly through time. Other subsamples of H. erectus (African, Chinese, and Indonesian) do not appear to increase significantly through time. Regression results generally corroborate results of the test for trend. Spatial and temporal variation may characterize evolution of cranial capacity in H. erectus. Different patterns of cranial capacity evolution may distinguish H. erectus from early H. sapiens.
Body size is one of the most important characteristics of any animal because it affects a range of behavioral, ecological, and physiological traits including energy requirements, choice of food, reproductive strategies, predation risk, range size, and locomotor style. This article focuses on the implications of being large bodied for Homo erectus females, estimated to have been over 50% heavier than average australopithecine females. The energy requirements of these hominins are modeled using data on activity patterns, body mass, and life history from living primates. Particular attention is given to the inferred energetic costs of reproduction for Homo erectus females based on chimpanzee and human reproductive scheduling. Daily energy requirements during gestation and lactation would have been significantly higher for Homo erectus females, as would total energetic cost per offspring if the australopithecines and Homo erectus had similar reproductive schedules (gestation and lactation lengths and interbirth intervals). Shortening the interbirth interval could considerably reduce the costs per offspring to Homo erectus and have the added advantage of increasing reproductive output. The mother would, however, incur additional daily costs of caring for the dependent offspring. If Homo erectus females adopted this reproductive strategy, it would necessarily imply a revolution in the way in which females obtained and utilized energy to support their increased energetic requirements. This transformation is likely to have occurred on several levels involving cooperative economic division of labor, locomotor energetics, menopause, organ size, and other physiological mechanisms for reducing the energetic load on females.
Analyses of the KNM-WT 15000 Homo erectus juvenile male partial skeleton from Kenya concluded that this species had a tall thin body shape due to specialized locomotor and climatic adaptations. Moreover, it was concluded that H. erectus pelves were obstetrically restricted to birthing a small-brained altricial neonate. Here we describe a nearly complete early Pleistocene adult female H. erectus pelvis from the Busidima Formation of Gona, Afar, Ethiopia. This obstetrically capacious pelvis demonstrates that pelvic shape in H. erectus was evolving in response to increasing fetal brain size. This pelvis indicates that neither adaptations to tropical environments nor endurance running were primary selective factors in determining pelvis morphology in H. erectus during the early Pleistocene.
Homo erectus inhabited a wide geographic area of Asia, ranging from 40 degrees north latitude in China to 8 degrees south latitude in island Southeast Asia. Yet variation within Asian H. erectus and its relation to ecological and temporal parameters have been little studied. I synthesize the revised radiometric chronologies for hominid sites in Asia and their relation to new oxygen isotope curves (proxies for climatic fluctuations and landbridge connections). These data suggest substantial opportunities in the later Pleistocene for both regional isolation and gene flow between hominids in mainland and Southeast Asia. They also suggest that the most northerly located Chinese sites (Zhoukoudian and Nanjing) may have been occupied during sequential, interglacial periods. Probably reflecting these periods of isolation, nonmetric features and principal components analysis (PCA) of calvarial shape suggest regional differentiation between northern Asian and Southeast Asian H. erectus. The most recent Southeast Asian fossils (e.g., Ngandong) conform to the Southeast Asian pattern. Except perhaps in brain size, there is no evidence that the temporally intermediate Chinese fossils are intermediate in morphology between older and younger Indonesian fossils. In fact, northern Chinese calvaria are easier to exclude from the larger Asian H. erectus hypodigm than are the Ngandong fossils. The Chinese specimens differ from the others based on their narrower occipitals and frontals for their cranial size. The Chinese sample from Zhoukoudian alone is thus not a good proxy for the morphology and variation seen within Asian H. erectus. Both the Chinese and late Indonesian samples exhibit less variation than does the early Indonesian sample; this along with their shared morphological bauplan suggests a common origin and no more than subspecific differentiation. This shared morphology, despite regional differences, was likely maintained by the increasing intensity of multiple glaciations (and longer-lasting land bridge connections) between mainland and island Southeast Asia during the last million years.
The first discoveries of Homo erectus were made in Java late in the 19th century. Since then many fossils have come to light in Africa as well as Asia. Homo erectus seems to have evolved in Africa before spreading to other regions of the Old World. This occurred over a long period of time, during which the populations changed relatively little. Only towards the close of the middle Pleistocene are there signs of change in evolutionary tempo leading to the appearance of more advanced humans. This book provides a wealth of information about individual crania, jaws and postcranial remains and will serve as an important guide to the anatomy of Homo erectus. It also documents the history of this extinct human species and suggests a route whereby Homo erectus may have given rise to people more like Homo sapiens.
The worldwide association of H. erectus with elephants is well documented and so is the preference of humans for fat as a source of energy. We show that rather than a matter of preference, H. erectus in the Levant was dependent on both elephants and fat for his survival. The disappearance of elephants from the Levant some 400 kyr ago coincides with the appearance of a new and innovative local cultural complex--the Levantine Acheulo-Yabrudian and, as is evident from teeth recently found in the Acheulo-Yabrudian 400-200 kyr site of Qesem Cave, the replacement of H. erectus by a new hominin. We employ a bio-energetic model to present a hypothesis that the disappearance of the elephants, which created a need to hunt an increased number of smaller and faster animals while maintaining an adequate fat content in the diet, was the evolutionary drive behind the emergence of the lighter, more agile, and cognitively capable hominins. Qesem Cave thus provides a rare opportunity to study the mechanisms that underlie the emergence of our post-erectus ancestors, the fat hunters.
Preface 1. Introduction 2. Homo erectus in the Far East 3. Homo erectus at Olduvai Gorge 4. Discoveries from the Turkana basin and other localities in sub-Saharan Africa 5. Northwest Africa 6. Comparisons of African hominids with Asian Homo erectus 7. Homo erectus as a paleospecies 8. The transition to more modern forms 9. Summary and prospects for further research References Author index Subject index.
Since its discovery within a travertine quarry, the fragmentary cranium of the only known Turkish Homo erectus, the Kocabaş hominid, has led to conflicting biochronological estimations. First estimated to be ∼500ka old, the partial skull presents a combination of archaic and evolved features that puts it as an intermediate specimen between the Dmanisi fossils (Homo georgicus) and the Chinese Zhoukoudian skulls (Homo erectus) respectively dated to 1.8 to ∼0.8Ma. Here we present a multidisciplinary study combining sedimentological, paleontological and paleoanthropological observations together with cosmogenic nuclide concentration and paleomagnetic measurements to provide an absolute chronological framework for the Upper fossiliferous Travertine unit where the Kocabaş hominid and fauna were discovered. The 26Al/10Be burial ages determined on pebbles from conglomeratic levels framing the Upper fossiliferous Travertine unit, which exhibits an inverse polarity, constrains its deposition to before the Cobb Mountain sub-chron, that is between 1.22 and ∼1.5Ma. The alternative match of the normal polarity recorded above the travertine with the Jaramillo subchron (lower limit 1.07 Ma) may also be marginally compatible with cosmogenic nuclides interpretation, thus the proposed minimum age of 1.1 Ma for the end of massive travertine deposition. The actual age of the fossils is likely to be in the 1.1–1.3 Ma range. This absolute date is in close agreement with the paleoanthropological conclusions based on morphometric comparisons implying that Kocabaş hominid belongs to the Homo erectus s.l. group that includes Chinese and African fossils, and is different from Middle and Upper Pleistocene specimens. Furthermore, this date is confirmed by the large mammal assemblage, typical of the late Villafranchian. Because it attests to the antiquity of human occupation of the Anatolian Peninsula and one of the waves of settlements out of Africa, this work challenges the current knowledge of the Homo erectus dispersal over Eurasia.
Bipedalism is a defining feature of the human lineage. Despite evidence that walking on two feet dates back 6-7 Ma, reconstructing hominin gait evolution is complicated by a sparse fossil record and challenges in inferring biomechanical patterns from isolated and fragmentary bones. Similarly, patterns of social behavior that distinguish modern humans from other living primates likely played significant roles in our evolution, but it is exceedingly difficult to understand the social behaviors of fossil hominins directly from fossil data. Footprints preserve direct records of gait biomechanics and behavior but they have been rare in the early human fossil record. Here we present analyses of an unprecedented discovery of 1.5-million-year-old footprint assemblages, produced by 20+ Homo erectus individuals. These footprints provide the oldest direct evidence for modern human-like weight transfer and confirm the presence of an energy-saving longitudinally arched foot in H. erectus. Further, print size analyses suggest that these H. erectus individuals lived and moved in cooperative multi-male groups, offering direct evidence consistent with human-like social behaviors in H. erectus.
It has been argued that Homo erectus is a species confined to Asia. Specialized characters displayed by the Indonesian and Chinese skulls are said to be absent in material from eastern Africa, and individuals from Koobi Fora and Nariokotome are now referred by some workers to H. ergaster. This second species is held to be the ancestor from which later human populations are derived. The claim for two taxa is evaluated here with special reference to the facial skeleton. Asian fossils examined include Sangiran 4 and Sangiran 17, several of the Ngandong crania, Gongwangling, and of course the material from Zhoukoudian described by Weidenreich ([1943] Palaeontol. Sin. [New Ser. D] 10:1-484). African specimens compared are KNM-ER 3733 and KNM-ER 3883 from Koobi Fora and KNM-WT 15000 from Nariokotome. Hominid 9 from Olduvai is useful only insofar as the brows and interorbital pillar are preserved. Neither detailed anatomical comparisons nor measurements bring to light any consistent patterns in facial morphology which set the African hominids apart from Asian H. erectus. Faces of the African individuals do tend to be high and less broad across the orbits. Both of the Koobi Fora crania but not KNM-WT 15000 have nasal bones that are narrow superiorly, while the piriform aperture is relatively wide. In many other characters, including contour of the supraorbital torus, glabellar prominence, nasal bridge dimensions, internasal keeling, anatomy of the nasal sill and floor, development of the canine jugum, orientation of the zygomaticoalveolar pillar, rounding of the anterolateral surface of the cheek, formation of a malar tubercle, and palatal rugosity, there is variation among individuals from localities within the major geographic provinces. Here it is not possible to identify features that are unique to either the Asian or African assemblages. Additional traits such as a forward sloping "crista nasalis," presence of a "sulcus maxillaris," a high (and massive) cheek coupled with some flexion of the malar pillar, and a posterior position for the incisive canal are present in all groups. These characters seem to be plesiomorphic, in comparison to the derived states evolved in later humans. Much or all of the variation in facial form can be attributed to sex dimorphism and/or local differentiation of populations within the Asian and African geographic regions. Metric differences among the fossils are comparable to those documented in a subset of recent H. sapiens, and there is no evidence that the Pleistocene specimens show greater dispersion than expected within a single species. This finding is generally in keeping with observations made on other parts of the cranium, lower jaw, and teeth. All of the hominids can be placed in H. erectus. Although its phylogenetic origins remain obscure, this lineage must be rooted in Africa. The species flourished for a long time. At several sites in China, H. erectus is known from deposits of the later Middle Pleistocene, while at Ngandong in Indonesia, archaic people may have survived even into the Late Pleistocene (Swisher et al. [1996] Science 274:1870-1874). The Ngandong fossils may record the last appearance of the lineage.
Abstract The state of information bearing on Homo erectus as developed since about 1960 is surveyed, with the resulting effects on problems. Definitions of H. erectus still rest on the Far Eastern samples (Chou‐k'ou‐tien/Java), and thus relate to late Lower to middle Middle Pleistocene material. Numerous important individual finds, however, have expanded the total: extension of the early and very early Sangiran material; very early to later in Africa, and relatively late in Europe. Datings remain uncertain or controversial within broad limits, but with some important successes and revisions. Discussion by authors of problems concerns degree of divergence among H. erectus populations and rate of evolutionary change; both appear relatively slight, but the data are inadequate for much present judgment. The apparent zone of transition to more advanced morphology ( H. sapiens , sensu lato) by the late Middle Pleistocene better reflects signs of regional divergence. Some writers—not all—believe that even the earliest European fossils known (e.g., Petralona) had already advanced to a H. sapiens basic level, with later change in the direction of Neanderthals. A separate African phylum, from OH 9, is also suggested; recent Chinese finds may provide a third different post‐ erectus population before the Upper Pleistocene. Taxonomic expression of all this gives some problems.
Hominid fossils from Ngandong and Sambungmacan, Central Java, are considered the most morphologically advanced representatives of Homo erectus. Electron spin resonance (ESR) and mass spectrometric U-series dating of fossil bovid teeth collected from the hominid-bearing levels at these sites gave mean ages of 27 +/- 2 to 53.3 +/- 4 thousand years ago; the range in ages reflects uncertainties in uranium migration histories. These ages are 20,000 to 400,000 years younger than previous age estimates for these hominids and indicate that H. erectus may have survived on Java at least 250,000 years longer than on the Asian mainland, and perhaps 1 million years longer than in Africa. The new ages raise the possibility that H. erectus overlapped in time with anatomically modern humans (H. sapiens) in Southeast Asia.
Homo erectus was the first human lineage to disperse widely throughout the Old World, the only hominin in Asia through much of the Pleistocene, and was likely ancestral to H. sapiens. The demise of this taxon remains obscure because of uncertainties regarding the geological age of its youngest populations. In 1996, some of us co-published electron spin resonance (ESR) and uranium series (U-series) results indicating an age as young as 35-50 ka for the late H. erectus sites of Ngandong and Sambungmacan and the faunal site of Jigar (Indonesia). If correct, these ages favor an African origin for recent humans who would overlap with H. erectus in time and space. Here, we report (40)Ar/(39)Ar incremental heating analyses and new ESR/U-series age estimates from the "20 m terrace" at Ngandong and Jigar. Both data sets are internally consistent and provide no evidence for reworking, yet they are inconsistent with one another. The (40)Ar/(39)Ar analyses give an average age of 546±12 ka (sd±5 se) for both sites, the first reliable radiometric indications of a middle Pleistocene component for the terrace. Given the technical accuracy and consistency of the analyses, the argon ages represent either the actual age or the maximum age for the terrace and are significantly older than previous estimates. Most of the ESR/U-series results are older as well, but the oldest that meets all modeling criteria is 143 ka+20/-17. Most samples indicated leaching of uranium and likely represent either the actual or the minimum age of the terrace. Given known sources of error, the U-series results could be consistent with a middle Pleistocene age. However, the ESR and (40)Ar/(39)Ar ages preclude one another. Regardless, the age of the sites and hominins is at least bracketed between these estimates and is older than currently accepted.
The Sambungmacan (Sm) 3 calvaria, discovered on Java in 1977, was illegally removed from Indonesia in 1998 and appeared in New York City in early 1999 at the Maxilla & Mandible, Ltd. natural history shop. Here we undertake an analysis of its phylogenetic and systematic position using geometric morphometrics and comparative morphology. The coordinates of points in the sagittal plane from glabella to opisthion were resampled to yield "lines" of 50 semi-landmarks. Coordinates of glabella, bregma, lambda, inion, and opisthion were also collected and analyzed separately. Casts of Homo erectus fossils from Indonesia, China, and Kenya and of "archaic H. sapiens" from Kabwe and Petralona, as well as 10 modern human crania, were used as the primary comparative sample. The modern humans were well separated from the fossils in a graphical superimposition of Procrustes-aligned semi-landmarks as well as in principal component and canonical discriminant analyses. In all of these, Sm 3 falls intermediate between the fossil and modern groups. Morphological comparisons of Sm 3 with a selection of Homo erectus fossils revealed its greatest similarity to specimens from Ngandong and the Sm 1 calvaria. Compared to all other H. erectus, Sm 3 was distinctive in its more vertical supratoral plane, less anteriorly projecting glabella and less sharply angled occiput. In these features it was somewhat similar to modern humans. It is not yet possible to determine if this similarity implies an evolutionary relationship or (more likely) individual or local populational variation. Several features of Sm 3 (small size, gracile supraorbital torus and lack of angular torus, and position in principal component analysis) suggest that it was a female. The use of geometric morphometrics provides a means to statistically test the shapes of such fossils in a manner not easily duplicated by other methods. The intermediate position of Sm 3 between fossil and modern samples in several different subanalyses exemplifies the value of this approach.
Bycatch studies have largely ignored population level effects on fish species of little commercial interest. Here we analyze bycatch of the lined seahorse (Hippocampus erectus) in the bait-shrimp trawl fishery in Hernando Beach, Florida, providing the first fisheries data for this species. Based on catch per unit of effort (CPUE), size, sex, and reproductive status of trawled H. erectus, 1) approximately 72,000 seahorses were caught annually by this fleet, from a population of unknown size, 2) trawling affected population cohorts differentially because of temporal and spatial variation in CPUE and population size, and 3) a greater proportion of females than males was removed in trawling. Our findings suggest that trawling may affect seahorse populations through direct mortality, social disruption, and habitat damage. However, the lack of specific abundance or catchability estimates for H. erectus means that the precise impact of trawling on this fish remains uncertain. This paper focuses attention on the need for research and monitoring of small fishes that are caught incidentally in nonselective gear.
Abstract New brain endocast reconstructions of Homo erectus discoveries from Indonesia since 1963 ( H. erectus VI, 1963; VII, 1965; VIII, 1969) have been made and their volumes determined. In addition, older discoveries ( H. erectus I, 1891; II, 1937; IV, 1937–38) have been reendocast and reconstructed, and have yielded volumes considerably different from those previously published. This is particularly so in the case of Dubois's original discovery, which yields a volume of 940 ml rather than the widely quoted volume of 750 ml. In addition, a number of morphological observations regarding hemispheric asymmetries (petalias) are provided, which suggest a condition similar to modern Homo sapiens .
Tham Khuyen Cave (Lang Son Province, northern Vietnam) is one of the more significant sites to yield fossil vertebrates in east Asia. During the mid-1960s, excavation in a suite of deposits produced important hominoid dental remains of middle Pleistocene age. We undertake more rigorous analyses of these sediments to understand the fluvial dynamics of Pleistocene cave infilling as they determine how skeletal elements accumulate within Tham Khuyen and other east Asian sites. Uranium/thorium series analysis of speleothems brackets the Pleistocene chronology for breaching, infilling, and exhuming the regional paleokarst. Clast analysis indicates sedimentary constituents, including hominoid teeth and cranial fragments accumulated from very short distances and under low fluvial energy. Electron spin resonance analysis of vertebrate tooth enamel and sediments shows that the main fossil-bearing suite (S1-S3) was deposited about 475 thousand years ago. Among the hominoid teeth excavated from S1-S3, some represent Homo erectus and Gigantopithecus blacki. Criteria are defined to differentiate these teeth from more numerous Pongo pygmaeus elements. The dated co-occurrence of Homo erectus and Gigantopithecus blacki at Tham Khuyen helps to establish the long co-existence of these two species throughout east Asia during the Early and Middle Pleistocene.
The evidence presented deals not with the early states of human evolution, but with the unequivocal occurrence of H. erectus from the Koobi Fora Formation, east of Lake Turkana. This chapter presents decisive evidence that shows the existence of two contemporaneous hominid species in the Koobi Fora area. The cranium consists of a complete calvaria and a great deal of the facial skeleton, including the nasal and zygomatic bones. The contemporaneity of Homo erectus and a robust Australopithecus is now clearly established over the period during which the Upper Member of the Koobi Fora Formation was deposited. The single species hypothesis has served a useful purpose in focusing attention on variability among the early hominids and also on the ecological consequences of hominid adaptations. Among the variety of hypotheses put forward to accommodate the evidence in an evolutionary framework, the most explicit and directly simple is the single species hypothesis.