Nacurrie 1: Mark of ancient Java, or a caring mother’s hands, in terminal Pleistocene Australia?

There has been a protracted debate over the evidence for intentional cranial modification in the terminal Pleistocene Australian crania from Kow Swamp and Coobool Creek. Resolution of this debate is crucial to interpretations of the significance of morphological variation within terminal Pleistocene-early Holocene Australian skeletal materials and claims of a regional evolutionary sequence linking Javan Homo erectus and Australian Homo sapiens. However, morphological comparisons of terminal Pleistocene and recent Australian crania are complicated by the significantly greater average body mass in the former. Raw and size-adjusted metric comparisons of the terminal Pleistocene skeleton from Nacurrie, south-eastern Australia, with modified and unmodified H. sapiens and H. erectus, identified a suite of traits in the frontal, parietal, and occipital bones associated with intentional modification of a neonate’s skull. These traits are also present in some of the crania from Kow Swamp and Coobool Creek, which are in close geographic proximity to Nacurrie, but not in unmodified H. sapiens or Javan H. erectus. Frontal bone morphology in H. erectus was distinct from all of the Australian H. sapiens samples. During the first six months of life, Nacurrie’s vault may have been shaped by his mother’s hands, rather than though the application of fixed bandages. Whether this behaviour persisted only for several generations, or hundreds of years, remains unknown. The reasons behind the shaping of Nacurrie’s head, aesthetics or otherwise, and why this cultural practice was adopted and subsequently discontinued, will always remain a matter of speculation.     

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.     

Human talus bones from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca,Burgos, Spain)

Here we present and describe comparatively 25 talus bones from the Middle Pleistocene site of the Sima de los Huesos (SH) (Sierra de Atapuerca, Burgos, Spain). These tali belong to 14 individuals (11 adult and three immature). Although variation among Middle and Late Pleistocene tali tends to be subtle, this study has identified unique morphological characteristics of the SH tali. They are vertically shorter than those of Late Pleistocene Homo sapiens, and show a shorter head and a broader lateral malleolar facet than all of the samples. Moreover, a few shared characters with Neanderthals are consistent with the hypothesis that the SH population and Neanderthals are sister groups. These shared characters are a broad lateral malleolar facet, a trochlear height intermediate between modern humans and Late Pleistocene H. sapiens, and a short middle calcaneal facet. It has been possible to propose sex assignment for the SH tali based on their size. Stature estimates based on these fossils give a mean stature of 174.4 cm for males and 161.9 cm for females, similar to that obtained based on the long bones from this same site.     

New 1.5 million-year-old Homo erectus maxilla from Sangiran (Central Java, Indonesia)

Sangiran (Solo Basin, Central Java, Indonesia) is the singular Homo erectus fossil locale for Early Pleistocene Southeast Asia. Sangiran is the source for more than 80 specimens in deposits with ⁴⁰Ar/³⁹Ar ages of 1.51-0.9 Ma. In April 2001, we recovered a H. erectus left maxilla fragment (preserving P³- M²) from the Sangiran site of Bapang. The find spot lies at the base of the Bapang Formation type section in cemented gravelly sands traditionally called the Grenzbank Zone. Two meters above the find spot, pumice hornblende has produced an ⁴⁰Ar/³⁹Ar age of 1.51 ± 0.08 Ma. With the addition of Bpg 2001.04, Sangiran now has five H. erectus maxillae. We compare the new maxilla with homologs representing Sangiran H. erectus, Zhoukoudian H. erectus, Western H. erectus (pooled African and Georgian specimens), and Homo habilis. Greatest contrast is with the Zhoukoudian maxillae, which appear to exhibit a derived pattern of premolar-molar relationships compared to Western and Sangiran H. erectus. The dental patterns suggest distinct demic origins for the earlier H. erectus populations represented at Sangiran and the later population represented at Zhoukoudian. These two east Asian populations, separated by 5000 km and nearly 800 k.yr., may have had separate origins from different African/west Eurasian populations.     

Dental microwear texture analysis of hominins recovered by the Olduvai Landscape Paleoanthropology Project, 1995-2007

Dental microwear analysis has proven to be a valuable tool for the reconstruction of aspects of diet in early hominins. That said, sample sizes for some groups are small, decreasing our confidence that results are representative of a given taxon and making it difficult to assess within-species variation. Here we present microwear texture data for several new specimens of Homo habilis and Paranthropus boisei from Olduvai Gorge, bringing sample sizes for these species in line with those published for most other early hominins. These data are added to those published to date, and microwear textures of the enlarged sample of H. habilis (n = 10) and P. boisei (n = 9) are compared with one another and with those of other early hominins. New results confirm that P. boisei does not have microwear patterns expected of a hard-object specialist. Further, the separate texture complexity analyses of early Homo species suggest that Homo erectus ate a broader range of foods, at least in terms of hardness, than did H. habilis, P. boisei, or the “gracile” australopiths studied. Finally, differences in scale of maximum complexity and perhaps textural fill volume between H. habilis and H. erectus are noted, suggesting further possible differences between these species in diet.     

Modern human origins: continuity, replacement, and masticatory robusticity in Australasia

Morphological evidence for a Multiregional (MR) model of human origins is suggested by a series of “linking traits” seen in the crania of late Javanese Homo erectus from Ngandong and anatomically modern Australian crania. A few studies that consider the genetic, structural, or functional aspects of these regional traits suggest their appearance is heavily influenced not by shared phylogeny but by a common “strong” masticatory pattern. Using dental occlusal areas, external mandibular metrics, internal biomechanical properties of the mandibular corpus measured from CT scans, and nonmetric traits associated with the attachment of masticatory muscles, we test the hypothesis that Australians exhibit evidence of a “strong” masticatory pattern. We use a mixed-sex comparative human sample (n = 415) that includes precontact Alaskans from Point Hope and the Aleutian Islands, Californians, Peruvians, an urban forensic sample, and the late Pleistocene Afalou-Taforalt sample. In comparison with recent humans known to exhibit such patterns, Australian mandibles show none of the expected changes related to producing and dissipating heavy occlusal loads. This is true regardless of whether external or internal mandibular dimensions are considered, albeit Australians show large occlusal areas and relatively large section modulus indices. Thus, a prime functional argument proposed for the origin of some Australian regional features is not supported by these data.     

Middle pleistocene humans from africa

Patterns of human evolution in the Middle Pleistocene remain poorly understood. There is general consensus that by the onset of this time period, populations ofHomo erectus were dispersed from Africa into Eurasia, including the Far East. In the western part of this range (perhaps in Africa),Homo erectus then produced a daughter lineage exhibiting more advanced characters of the face, braincase and cranial base. How this new species should be defined is currently debated. In my view, fossils from sites such as Bodo and Broken Hill in Africa may be lumped with material from earlier Middle Pleistocene localities in Europe. Such a taxon is appropriately namedHomo heidelbergensis. Whether the hypodigm should be extended to include fossils from China is another question. In any case, this group of hominids is plausibly ancestral to both the specialized Neanderthals of Europe and more modern humans of the later Middle Pleistocene.     

Three newHomo erectus mandibles from Java

There are now eleven manidublar pieces from the Lower and Middle Pleistocene of Java, all but one being from the Sangiran site. All of these have been assigned toHomo erectus by most workers, while others have suggested as many as four different hominoid taxa. Sangiran 21 (Mandible E), Sangiran 22 (Mandible F), and Sangiran 37 (Mandible G) are described here fully for the first time. Sangiran 21, 22, and 27 all come from the Upper Pucangan Formation and date approximately 1.2 Myr. The new mandibles are morphologically compatible with theH. erectus, crania from Java.     

Body size and body shape in early hominins - implications of the Gona Pelvis

Discovery of the first complete Early Pleistocene hominin pelvis, Gona BSN49/P27, attributed to Homo erectus, raises a number of issues regarding early hominin body size and shape variation. Here, acetabular breadth, femoral head breadth, and body mass calculated from femoral head breadth are compared in 37 early hominin (6.0-0.26 Ma) specimens, including BSN49/P27. Acetabular and estimated femoral head sizes in the Gona specimen fall close to the means for non-Homo specimens (Orrorin tugenesis, Australopithecus africanus, Paranthropus robustus), and well below the ranges of all previously described Early and Middle Pleistocene Homo specimens. The Gona specimen has an estimated body mass of 33.2 kg, close to the mean for the non-Homo sample (34.1 kg, range 24-51.5 kg, n = 19) and far outside the range for any previously known Homo specimen (mean = 70.5 kg; range 52-82 kg, n = 17). Inclusion of the Gona specimen within H. erectus increases inferred sexual dimorphism in body mass in this taxon to a level greater than that observed here for any other hominin taxon, and increases variation in body mass within H. erectus females to a level much greater than that observed for any living primate species. This raises questions regarding the taxonomic attribution of the Gona specimen. When considered within the context of overall variation in body breadth among early hominins, the mediolaterally very wide Gona pelvis fits within the distribution of other lower latitude Early and Middle Pleistocene specimens, and below that of higher latitude specimens. Thus, ecogeographic variation in body breadth was present among earlier hominins as it is in living humans. The increased M-L pelvic breadth in all earlier hominins relative to modern humans is related to an increase in ellipticity of the birth canal, possibly as a result of a non-rotational birth mechanism that was common to both australopithecines and archaic Homo.     

Reconstructing cranial evolution in an extinct hominin

Homo erectus is the first hominin species with a truly cosmopolitan distribution and resembles recent humans in its broad spatial distribution. The microevolutionary events associated with dispersal and local adaptation may have produced similar population structure in both species. Understanding the evolutionary population dynamics of H. erectus has larger implications for the emergence of later Homo lineages in the Middle Pleistocene. Quantitative genetics models provide a means of interrogating aspects of long-standing H. erectus population history narratives. For the current study, cranial fossils were sorted into six major palaeodemes from sites across Africa and Asia spanning 1.8–0.1 Ma. Three-dimensional shape data from the occipital and frontal bones were used to compare intraspecific variation and test evolutionary hypotheses. Results indicate that H. erectus had higher individual and group variation than Homo sapiens, probably reflecting different levels of genetic diversity and population history in these spatially disperse species. This study also revealed distinct evolutionary histories for frontal and occipital bone shape in H. erectus, with a larger role for natural selection in the former. One scenario consistent with these findings is climate-driven facial adaptation in H. erectus, which is reflected in the frontal bone through integration with the orbits.     

Mandibular molar root morphology in Neanderthals and Late Pleistocene and recent Homo sapiens

Neanderthals have a distinctive suite of dental features, including large anterior crown and root dimensions and molars with enlarged pulp cavities. Yet, there is little known about variation in molar root morphology in Neanderthals and other recent and fossil members of Homo. Here, we provide the first comprehensive metric analysis of permanent mandibular molar root morphology in Middle and Late Pleistocene Homo neanderthalensis, and Late Pleistocene (Aterian) and recent Homo sapiens. We specifically address the question of whether root form can be used to distinguish between these groups and assess whether any variation in root form can be related to differences in tooth function. We apply a microtomographic imaging approach to visualise and quantify the external and internal dental morphologies of both isolated molars and molars embedded in the mandible (n = 127). Univariate and multivariate analyses reveal both similarities (root length and pulp volume) and differences (occurrence of pyramidal roots and dental tissue volume proportion) in molar root morphology among penecontemporaneous Neanderthals and Aterian H. sapiens. In contrast, the molars of recent H. sapiens are markedly smaller than both Pleistocene H. sapiens and Neanderthals, but share with the former the dentine volume reduction and a smaller root-to-crown volume compared with Neanderthals. Furthermore, we found the first molar to have the largest average root surface area in recent H. sapiens and Neanderthals, although in the latter the difference between M₁ and M₂ is small. In contrast, Aterian H. sapiens root surface areas peak at M₂. Since root surface area is linked to masticatory function, this suggests a distinct occlusal loading regime in Neanderthals compared with both recent and Pleistocene H. sapiens.     

“Meganthropus” cranial fossils from Java

There are now twelve significant hominid cranial fossils from the Lower and Middle Pleistocene of Java, all but two being from the Sangiran site. Most of this material is well-known in the literature, but three skulls, possibly representing “Meganthropus” are here described in detail for the first time. Most scholars have assigned them all toHomo erectus, while others have suggested that they represent as many as four different hominoid taxa. The author argues that they represent two possible species of hominids. “Meganthropus” I, II, and III are more massive than any of the knownH. erectus specimens. They are also relatively higher vaulted, apparently smaller brained, and have unusually thick lower occipital planes. “Meganthropus” may represent a species that separated fromH. erectus upon its arrival to Java.     

Mojokerto revisited: Evidence for an intermediate pattern of brain growth in Homo erectus

Brain development in Homo erectus is a subject of great interest, and the infant calvaria from Mojokerto, Indonesia, has featured prominently in these debates. Some researchers have suggested that the pattern of brain development in H. erectus resembled that of non-human apes, while others argue for a more human-like growth pattern. In this study, we retested hypotheses regarding brain ontogeny in H. erectus using new methods (resampling), and data from additional H. erectus crania. Our results reveal that humans achieve 62% (±10%) and chimpanzees 80% (±9%) of their adult endocranial volume by 0.5–1.5 years of age. Using brain mass data, humans achieve on average 65% and chimpanzees 81% of adult size by 0.5–1.5 years. When compared with adult H. erectus crania (n = 9) from Indonesian sites greater than 1.2 million years old, Mojokerto had reached ∼70% of its adult cranial capacity. Mojokerto thus falls almost directly between the average growth in humans and chimpanzees, and well within the range of both. We therefore suggest that brain development in H. erectus cannot be dichotomized as either ape-like or human-like; it was H. erectus-like. These data indicate that H. erectus may have had a unique developmental pattern that should be considered as an important step along the continuum of brain ontogeny between apes and humans.     

Taxonomic affinity of the early Homo cranium from Swartkrans,South Africa

A quantitative analysis that employs randomization methods and distance statistics has been undertaken in an attempt to clarify the taxonomic affinities of the partial Homo cranium (SK 847) from Member 1 of the Swartkrans Formation. Although SK 847 has been argued to represent early H. erectus, exact randomization tests reveal that the magnitude of differences between it and two crania that have been attributed to that taxon (KNM-ER 3733 and KNM-WT 15000) is highly unlikely to be encountered in a modern human sample drawn from eastern and southern Africa. Some of the variables that differentiate SK 847 from the two early H. erectus crania (e. g., nasal breadth, frontal breadth, mastoid process size) have been considered to be relevant characters in the definition of that taxon. Just as the significant differences between SK 847 and the two early H. erectus crania make attribution of the Swartkrans specimen to that taxon unlikely, the linkage of SK 847 to KNM-ER 1813, and especially Stw 53, suggests that the Swartkrans cranium may have its closest affinity with H. habilis sensu lato. Differences from KNM-ER 1813, however, hint that the South African fossils may represent a species of early Homo that has not been sampled in the Plio-Pleistocene of eastern Africa. The similarity of SK 847 and Stw 53 may support faunal evidence which suggests that Sterkfontein Member 5 and Swartkrans Member 1 are of similar geochronological age. © 1993 Wiley-Liss, Inc.     

Where are inion and endinion? Variations of the exo- and endocranial morphology of the occipital bone during hominin evolution

The occipital bone is frequently investigated in paleoanthropological studies because it has several features that help to differentiate various fossil hominin species. Among these features is the separation between inion and endinion, which has been proposed to be an autapomorphic trait in (Asian) Homo erectus. Methodologies are developed here to quantify for the first time the location of these anatomical points, and to interpret their variation due to the complex interactions between exocranial and endocranial size and shape of the occipital and nuchal planes, as well as the occipital lobes and cerebellum. On the basis of our analysis, neither ‘the separation between inion and endinion’ nor ‘endinion below inion’ can be considered as an autapomorphic trait in H. erectus, since this feature is a condition shared by extant African great apes and fossil hominins. Moreover, our results show that the exo- and endocranial anatomy of the occipital bone differs between hominins (except Paranthropus boisei specimens and KNM-ER 1805) and great apes. For example, chimpanzees and bonobos are characterized by a very high position of inion and their occipital bone shows an antero-posterior compression. However, these features are partly correlated with their small size when compared with hominins. Asian H. erectus specimens have a thick occipital torus, but do not differ from other robust specimens, neither in this feature nor in the analysed exo- and endocranial proportions of the occipital bone. Finally, the apparent brain size reduction during the Late Pleistocene and variation between the sexes in anatomically modern humans (AMH) reflect that specimens with smaller brains have a relatively larger posterior height of the cerebellum. However, this trend is not the sole explanation for the ‘vertical shift’ of endinion above inion that appears occasionally and exclusively in AMH.     

Cranial remains of Homo erectus from Beds II and IV,Olduvai Gorge,Tanzania

Cranial remains of hominids 9 and 12 from Olduvai Gorge are described in detail. O.H. 9 consists of a heavily built braincase, partly damaged and lacking the face, while O.H. 12 is less complete. The Bed II specimen is about 1.2 million years in age and shows anatomical similarities to the cranium designated ER-3733 from Koobi Fora, east of Lake Turkana. Together these African fossils provide valuable information about Homo erectus in the later Lower Pleistocene. Comparisons of O.H. 9 with several of the Choukoutien crania are also carried out. These Chinese and other Asian remains of Homo erectus cannot be placed in a secure chronological framework, but all of the material should be studied systematically in order to assess relatedness among what must be several different populations.     

Les restes humains du Pliocène final et du début du Pléistocène inférieur de Dmanissi, Géorgie (1991-2000). I - Les crânes, D 2280, D 2282, D 2700

Since 1991, several human remains: 5 skulls, 4 mandibles and numerous postcranial fragments have been discovered on the Dmanissi prehistoric open site. It is an exceptional discovery due to the stratigraphical, paleontological and cultural context, which is well known and accurately well dated (Upper Pliocene-Early Pleistocene). Most of the hominids discovered in the level V and VI are dated between 1.81 My (level V) and 1.77 My (level VI) corresponding to a 40,000 years period. The assemblage of fossil human remains is peculiar due to (1) the quality of bone representation (distinct parts of the skeleton are preserved: skull, thorax, upper and lower limbs, belt), (2) the high degree of bone preservation (skulls and long bones are entire, rarely broken or crushed), (3) the diversity age at death estimated for each of the 5 individuals (3 adults, 1 young adult, 1 adolescent of both sexes). The study dealing with the first discovered mandibles and skulls has begun with Leo Gabounia since 1991 and represents several interests: 1) a paleoanthropological interest: the Dmanissi skulls are characterized by their small size; they are short, narrow and low. The skullcaps are less elevated than those of the Homo erectus group and even those of Homo ergaster. They are more elevated than those of Homo habilis and very close to Homo rudolfensis. The elevation and the transversal development of the middle part of the skull in the parietotemporal region are more significant: the Dmanissi specimens are intermediate between Homo habilis and Homo ergaster. In term of cranial capacity, a similar trend is observed. Generally speaking, the skull is slender. The vault is more flat than in Homo erectus, the frontal bone is less developed, divergent and the postorbital constriction is strong. The temporal bone is long, flat and the mastoid part is short. The upper part of the occipital bone is low and narrow. Crests are thin, less developed than in the Homo erectus group. The superior temporal crests are in a high position and a torus angularis is present on the adult-male specimen. The glenoid cavity is large with strong edges. The petrotympanic region is slender with a tympanic circle individualized and it shows a horizontal rotation in a posterior position, which is distinct from Homo erectus. The orthognathic trend of the face distinguishes the Dmanissi specimens from the early Pleistocene hominids (Homo habilis, Homo ergaster) and from the first Eurasian Homo erectus. Nevertheless, the subnasal region of the face is projected. The morphology of the mid-face, showing a developed pillar of the canine, an inframalar incurvation and an anterior position of the root of the zygomaticomaxillary crest, suggests strong masticatory stress. Considering the overall morphology, cranial and metrical features, the Dmanissi fossil skulls are intermediate to the Homo habilis-rudolfensis group and Homo ergaster while they are closer to the former and peculiarly to Homo rudolfensis (ER 1470). However, the Dmanissi fossil skulls are distinct from Homo rudolfensis by numerous features and among them: by their large maximum cranial width (Euryon-Euryon), the posterior rotation of their petrotympanic structure and the strong development of the pillar of their canine. Due to the gracility of their face, the narrowness of their occipital bone, and their cranial base pattern (mastoid region and petrotympanic structure), the Dmanissi fossil skulls are different from the Homo erectus group: 2) the abundance of the human fossils discovered in Dmanissi site provides information about the biodiversity of these hominids with the establishment of the morphological features related to either growth or sexual patterns: 3) compared to modern humans, the Dmanissi fossil skulls seem to follow a different growth pattern. The present study of the fossil skulls discovered is a pioneer step. Indeed, the Dmanissi site has yielded the oldest evidences of the first settlements in Eurasia, which were, until now, attributed to Homo erectus. The Dmanissi fossil skulls are close to the Homo habilis-rudolfensis African group. We attribute these hominids to Homo georgicus.