Metatarsal robusticity in primates and a few other plantigrade mammals

By using a new technique for determining relative metatarsal robusticity, the distribution of the 1+5 pattern (in which M5 is the second most robust metatarsal after M1) and 15 pattern (in which M5 is the least robust of all metatarsals) was established in primates and a few other plantigrade mammals. The first pattern is associated with a terrestrial and the second with an arboreal substrate. Robusticity formulae are not connected with specific locomotor patterns, but a total robusticity quotient is associated with these patterns and substrate preference as well. Changes in substrate preference are accompanied by changes of total robusticity, an increased number of permutations and ultimately a change of the robusticity pattern. Intermetatarsal robusticity gradients are related to the direction and intensity of muscular activity. A combined analysis of all factors can reveal a great deal of the locomotor history of a taxon.Also ofThe Institute of Applied Biology, New York.     

Implications of relative robusticity in the Olduvai metatarsus

The relative robusticity of the metatarsals in OH-8 is reviewed in light of the normal variation in metatarsal robusticity in Pan and Homo. The pattern in OH-8 is found to be fully commensurate with a striding gait.     

Postcranial robusticity in Homo. I: Temporal trends and mechanical interpretation

Temporal trends in postcranial robusticity within the genus Homo are explored by comparing cross-sectional diaphyseal and articular properties of the femur, and to a more limited extent, the humerus, in samples of Recent and earlier Homo. Using both theoretical mechanical models and empirical observations within Recent humans, scaling relationships between structural properties and bone length are developed. The influence of body shape on these relationships is considered. These scaling factors are then used to standardize structural properties for comparisons with pre-Recent Homo (Homo sp. and H. erectus, archaic H. sapiens, and early modern H. sapiens). Results of the comparisons lead to the following conclusions: 1) There has been a consistent, exponentially increasing decline in diaphyseal robusticity within Homo that has continued from the early Pleistocene through living humans. Early modern H. sapiens are closer in shaft robusticity to archaic H. sapiens than they are to Recent humans. The increase in diaphyseal robusticity in earlier Homo is a result of both medullary contraction and periosteal expansion relative to Recent humans. 2) There has been no similar temporal decline in articular robusticity within Homo–relative femoral head size is similar in all groups and time periods. Thus, articular to shaft proportions are different in pre-Recent and Recent Homo. 3) These findings are most consistent with a mechanical explanation (declining mechanical loading of the postcranium), that acted primarily through developmental rather than genetic means. The environmental (behavioral) factors that brought about the decline in postcranial robusticity in Homo are ultimately linked to increases in brain size and cultural-technological advances, although changes in robusticity lag behind changes in cognitive capabilities. © 1993 Wiley-Liss, Inc.     

A reappraisal of variation in hominid mandibular corpus dimensions

The relationship between breadth and height of the mandibular corpus has been investigated in a sample of 77 hominid mandibles. An interspecific allometric increase in robusticity with size occurs between four taxonomic subgroups of Australopithecus, but subgroups of Homo vary in robusticity while differing little in size. Within taxonomic subgroups, variation in breadth is not significantly related to variation in height among the “gracile” australapithecines; however, it is isometrically related to height in the “robust” australopithecines and bears an allometric relationship to height in Homo. Thus, robusticity, in conjunction with size, may provide a useful indicator of the taxonomic affinities of hominid mandibles.     

Hand use and metacarpal robusticity in Catarrhini

Recent studies,Rightmire (1972) andDay andScheuer (1973), have investigated the affinities of early hominid metacarpals from Swartkrans. Because of its extensive use in the analysis of metatarsals, the index of robusticity has also been applied to some fossil metacarpals. From the metatarsal analyses, it has been shown that within a group a variety of robusticity patterns exist with the average pattern occuring approximately 50 per cent of the time. This, coupled with the fact that it has also been shown that within the order Primates the pattern is not necessarily diagnostic of any locomotor category, has led us to investigate the usefulness of such an index in assessing primate metacarpals. In this study, metacarpal robusticity patterns and the Total Robusticity Quotient are established for seven cercopithecoid genera and the results correlated with hand use. It is found that although patterns of robusticity are not diagnostic of locomotor categories, the TRQ relates well to hand use: low TRQ's are found in primates which use their hands to walk on a flat substrate, while high TRQ's are diagnostic of arm swinging animals. Primates with reduced thumb use in a precision grip and little manipulative use of the hand have higher TRQ's than those with a good precision grip. The model derived from our sample of Cercopithecoidea is tested with a hominoid sample of four genera and found to be similarly applicable.     

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.     

An Early Pleistocene human pedal phalanx from Swartkrans,SKX 16699, and the antiquity of the human lateral forefoot

In order to assess the antiquity of derived human lateral (lesser) toe morphology, the SKX 16699 Early Pleistocene pedal proximal phalanx from Swartkrans (South Africa) was compared to samples of pedal phalanges attributed to Pliocene/Pleistocene australopithecines, Homo naledi and Late Pleistocene Homo. In contrast to australopith lateral phalanges, the SKX 16699 phalanx exhibits an absolutely (and probably relatively) short length, limited plantar diaphyseal curvature, proximal-to-midshaft and mid-dorsoplantar flexor sheath insertions, and a marked proximodorsal orientation of the metatarsal facet. SKX 16699 is intermediate between the australopith phalanges and later Homo ones in its modest dorsal diaphyseal curvature and mid-dorsoplantar metatarsophalangeal collateral ligament insertion areas. Its diaphyseal robustness is similar to that of Homo phalanges, but overlaps the range of later australopith ones. This combination of features and the close morphological affinities of SKX 16699 to later Homo proximal pedal phalanges suggest the emergence of a distinctly human lateral forefoot by the initial Early Pleistocene.     

The skeletal correlates of behavioral modification in the laboratory mouse (Mus musculus)

The effects of behavioral modification on the skeletal morphology of the laboratory mouse (Mus musculus) are investigated. Climibing, with increrased prehensile use of the foot, is found to bring about significant changes in metatarsal and long bone morphology. Differences in metatarsal robusticity related to weight-bearing differences associated with different locomotor patterns are reported for a natural history setting for Peromyscus.     

Changes in metatarsal robusticity following experimental surgery

When a marginal metatarsal (M1 or M5) is amputated in neonatal rats, robusticity increases in all other metatarsals in both sexes. The robusticity increase is greatest in the metatarsal adjacent to the operation and declines in the direction away from it. The tripod structure of the rat's foot with its maximum robusticity of the marginal metatarsals is retained. When both marginal metatarsals are removed, this tripod arrangement is less clearly retained. When robusticity of all remaining metatarsals is averaged in a total robusticity quotient, total robusticity increase is greater in M1 than in M5 removal and greatest when both are removed. Total increase is greater in females than in males. Metatarsal robusticity increases result from retardation of longitudinal growth and increase of weight. The fact that some robusticity increases coincide with lesser length but not with increased weight suggests that in the process of robusticity increase longitudinal growth retardation precedes that of weight increase. Lesser length and greater weight of a long bone occurs also when hypofunction is produced in neonatal rats. However, in hypofunction the thickness of the compacta is increased leading to an obliteration of the medullary canal, while in the experimental metatarsals of this study, submitted to greater weight-bearing stresses, this canal is never reduced or obliterated.     

Skeletal evidence for precision gripping inCebus apella

We compared the thumb morphology ofCebus apella to that of several other primate species in order to determine whether robust thumbs are associated with tool-use. We found that thumb robusticity was greater forCebus apella than for all other represented nonhuman species exceptGorilla gorilla. Further, thumb robusticity inCebus apella was similar to that ofAustralopithecus afarensis but lesser than that of other represented hominids, including modern humans. We propose that precision gripping similar to that which occurs in tool-using context amongCebus probably occurred among Australopithecines prior to the emergence of sophisticated tool behavior amongHomo andParanthropus.     

Reevaluation of the lumbosacral region of Oreopithecus bambolii

Functional interpretations of the postcranium of the late Miocene ape Oreopithecus bambolii are controversial. The claim that Oreopithecus practiced habitual terrestrial bipedalism is partly based on restored postcranial remains originally recovered from Baccinello, Tuscany ( Köhler and Moyà-Solà, 1997). The lower lumbar vertebrae of BA#72 were cited as evidence that Oreopithecus exhibits features indicative of a lordotic lumbar spine, including dorsal wedging of the vertebral bodies and a caudally progressive increase in postzygapophyseal interfacet distance. Here, we demonstrate why the dorsal wedging index value obtained by Köhler and Moyà-Solà (1997) for the BA#72 last lumbar vertebra is questionable due to distortion in that region, present a more reliable way to measure postzygapophyseal interfacet distance, and include an additional metric (laminar width) with which to examine changes in the transverse dimensions of the neural arches. We also quantify the external morphology of the BA#72 proximal sacrum, which, despite well-documented links between sacral morphology and bipedal locomotion, and excellent preservation of the sacral prezygapophyses, first sacral vertebral body, and right ala, was not evaluated by Köhler and Moyà-Solà (1997). Measures of postzygapophyseal interfacet distance and laminar width on the penultimate and last lumbar vertebrae of BA#72 reveal a pattern encompassed within the range of living nonhuman hominoids and unlike that of modern humans, suggesting that Oreopithecus did not possess a lordotic lumbar spine. Results further show that the BA#72 sacrum exhibits relatively small prezygapophyseal articular facet surface areas and mediolaterally narrow alae compared with modern humans, indicating that the morphology of the Oreopithecus sacrum is incompatible with the functional demands of habitual bipedal stance and locomotion. The Oreopithecus lumbosacral region does not exhibit adaptations for habitual bipedal locomotion.     

Variations and asymmetries in regional brain surface in the genus Homo

Paleoneurology is an important field of research within human evolution studies. Variations in size and shape of an endocast help to differentiate among fossil hominin species whereas endocranial asymmetries are related to behavior and cognitive function. Here we analyse variations of the surface of the frontal, parieto-temporal and occipital lobes among different species of Homo, including 39 fossil hominins, ten fossil anatomically modern Homo sapiens and 100 endocasts of extant modern humans. We also test for the possible asymmetries of these features in a large sample of modern humans and observe individual particularities in the fossil specimens.This study contributes important new information about the brain evolution in the genus Homo. Our results show that the general pattern of surface asymmetry for the different regional brain surfaces in fossil species of Homo does not seem to be different from the pattern described in a large sample of anatomically modern H. sapiens, i.e., the right hemisphere has a larger surface than the left, as do the right frontal, the right parieto-temporal and the left occipital lobes compared with the contra-lateral side. It also appears that Asian Homo erectus specimens are discriminated from all other samples of Homo, including African and Georgian specimens that are also sometimes included in that taxon. The Asian fossils show a significantly smaller relative size of the parietal and temporal lobes. Neandertals and anatomically modern H. sapiens, who share the largest endocranial volume of all hominins, show differences when considering the relative contribution of the frontal, parieto-temporal and occipital lobes. These results illustrate an original variation in the pattern of brain organization in hominins independent of variations in total size. The globularization of the brain and the enlargement of the parietal lobes could be considered derived features observed uniquely in anatomically modern H. sapiens.     

Cranio-dental evidence of a hominin-like hyper-masticatory apparatus in Oreopithecus bambolii. Was the swamp ape a human ancestor?

The phylogeny of Oreopithecus bambolii has been controversial since Johannes Hurzeler first argued that the Late Miocene (Tortonian) primate was a fossil hominin. While most paleontologist currently exclude Oreopithecus from human ancestry, recent postcranial evidence of hominin-like bipedalism and power precision grips in Oreopithecus has rekindled interest in the fossil Italian hominoid.In this study, a comparative review of hominoid cranio-dental morphology indicates that Oreopithecus possessed a suite of hominin-like characteristics that were apparently functionally related to powerful folivorous mastication. Since the oreopithecine dentition exhibited exceptional adaptations for folivory relative to most other extant and extinct hominoids, the accessory development of a hominin-like hyper-masticatory cranio-mandibular apparatus to further enhance plant comminution and digestibility is not surprising. However, the combination of hominin-like locomotive, manual, and masticatory functional attributes appears to substantiate Hurzeler's designation of O. bambolii as a Late Miocene hominin. Additionally, an extensive compilation of hominoid cranio-dental and postcranial characteristics strongly supports a close phylogenetic relationship between Oreopithecus and the earliest African hominins Sahelanthropus and Australopithecus.The wetland paleoecology of the Tortonian island of Tuscany–Sardinia suggests that Oreopithecus was a specialized semiaquatic folivore who apparently waded bipedally into freshwater swamps to feed on aquatic plants. However, the extensive wear on the oreopithecine canines and incisors along with their manual precision grips may indicate that freshwater invertebrates were also included in their diets. Such specialized aquatic feeding behavior by these ancient Italian swamp apes seems to support Alister Hardy's hypothesis that human bipedalism and power precision grips were inherited from Late Miocene hominin ancestors who originally utilized such functional attributes for aquatic foraging in shallow water environments. Additionally, the remarkably hominin-like cranio-dental morphology of O. bambolii suggests that modern omnivorous humans probably inherited a significant number of their cranio-dental characteristics from these highly specialized aquatic plant eating hominins.     

First hominine remains from a ∼1.0 million year old bone bed at Cornelia-Uitzoek, Free State Province, South Africa

We report here on evidence of early Homo around 1.0 Ma (millions of years ago) in the central plains of southern Africa. The human material, a first upper molar, was discovered during the systematic excavation of a densely-packed bone bed in the basal part of the sedimentary sequence at the Cornelia-Uitzoek fossil vertebrate locality. We dated this sequence by palaeomagnetism and correlated the bone bed to the Jaramillo subchron, between 1.07 and 0.99 Ma. This makes the specimen the oldest southern African hominine remains outside the dolomitic karst landscapes of northern South Africa. Cornelia-Uitzoek is the type locality of the Cornelian Land Mammal Age. The fauna contains an archaic component, reflecting previous biogeographic links with East Africa, and a derived component, suggesting incipient southern endemism. The bone bed is considered to be the result of the bone collecting behaviour of a large predator, possibly spotted hyaenas. Acheulian artefacts are found in small numbers within the bone bed among the fossil vertebrates, reflecting the penecontemporaneous presence of people in the immediate vicinity of the occurrence. The hominine tooth was recovered from the central, deeper part of the bone bed. In size, it clusters with southern African early Homo and it is also morphologically similar. We propose that the early Homo specimen forms part of an archaic component in the fauna, in parallel with the other archaic faunal elements at Uitzoek. This supports an emergent pattern of archaic survivors in the southern landscape at this time, but also demonstrates the presence of early Homo in the central plains of southern Africa, beyond the dolomitic karst areas.     

Metatarsal robusticity in bipedal rats

All metatarsals have a significantly greater robusticity in the male than in the female rat. The robusticity formula of the rat's foot is 1 > 5 > 2 > 3 > 4. In bipedal rats that formula remains unchanged, but the robusticity of the metatarsals is increased especially in females. The tripod arrangement of the human foot with its particular robustness of the marginal metatarsals 1 and 5 and a strong calcaneum has been related to upright posture. The similar robusticity pattern in the rat's marginal metatarsals 1 and 5 raises the question of whether that part of the formula might not represent a more general plantigrade pattern.     

Carbon isotopic record of terrestrial ecosystems spanning the Late Miocene extinction of Oreopithecus bambolii, Baccinello Basin (Tuscany, Italy)

Oreopithecus bambolii is a Late Miocene hominoid with an extensive fossil record in the Baccinello Basin (Tuscany, Italy), and was the only western European hominoid to survive a major extinction event ca. 9.6 Ma (millions of years ago). Oreopithecus lived in the insular Tusco-Sardinian paleobioprovince, where it evolved many unique anatomical specializations that make it important for understanding the mechanisms and history of Late Miocene hominoid evolution. The eventual extinction of Oreopithecus and its associated fauna ca. 6.5 Ma has generally been attributed to interaction with species that arrived from continental Europe following tectonic collision of the Tusco-Sardinian province with mainland Italy, but palynological, paleontological, and sedimentological records indicate an environmental shift toward more variable climate across the extinction event.To explore the possibility of environmental change as a contributing factor in the extinction of Oreopithecus, we developed a stable carbon and oxygen isotope record from organic matter in paleosols from the Baccinello Basin. These data show very low temporal and spatial variability (indicating plant ecosystem stability through time and space) and provide no evidence for ecologically significant changes in floral composition spanning the extinction event, suggesting that environmental change was not an underlying cause for the extinction of Oreopithecus and its associated fauna. The carbon isotope values fall entirely within the range of isotopic variability for modern plants following the C₃ photosynthetic pathway (trees, shrubs, cool-season grasses), indicating that C₄ vegetation (warm-season grasses) was not an important component of biomass. When corrected for temporal variation in the carbon isotopic composition of atmospheric carbon dioxide, the paleosol carbon isotope values are consistent with predicted values based on modern plants and the Baccinello palynoflora, supporting the reliability of paleosol isotopic records as paleoecological proxies.     

Olduvai Hominin 8 foot pathology: A comparative study attempting a differential diagnosis

Olduvai Hominin (OH) 8, a 1.76 million year old left foot skeleton, has osteophytic lipping on the metatarsal bases, which when compared to a modern sample, may help paleoanthropologists determine whether the foot bones represent an injured subadult or an osteoarthritic adult. This study compares the OH 8 lipping pattern to those of 140 individual Amerindians comprising four different age classes to determine whether the OH 8 lipping is likely to be age-related osteoarthritis. OH 8 metatarsal lipping followed a pattern similar to that determined in the comparative sample to be age-related osteoarthritis. Similarities include metatarsal base lipping that is frequently located on the dorsal surface, metatarsal base lipping that is more severe on the lateral metatarsals compared to the medial metatarsals, and the presence of a pseudojoint between metatarsal 1 and metatarsal 2. The chance of finding an individual with osteoarthritis lipping increases from 3.45% in the age group 18–22 years to 55% in individuals over 35 years. The chance of finding a pseudojoint increases from 1.32% in non-osteoarthritic individuals to 15.15% in individuals with osteoarthritis. Results from this study indicate that the OH 8 foot bones are most likely from an adult and more likely to belong to Paranthropus boisei, the skull of which was found in the same excavations with OH 8, than to the juvenile Homo habilis holotype.     

Hand biomechanics during simulated stone tool use

Human radial digits have derived features compared with apes, with long robust thumbs, relatively larger joint surfaces, and hypertrophic thenar muscles. Here we test the hypothesis that these features evolved in the context of making and using stone tools, specifically for producing large gripping forces and for countering large joint contact stresses. We used portable force plates simulating early stone tools to: 1) document and compare the magnitude of external/internal forces and joint stresses in the radial digits during hardhammer percussion and flake use, and 2) examine how variation in digit morphology affects muscle and joint mechanics during stone tool use. Force and kinematic data were collected from a sample representing normal variation in digit morphology (n = 25). The effects of digit size/shape on digit biomechanics were evaluated using partial correlations, controlling for tool reaction forces and impact velocities. Results show that individuals with longer digits require relatively less muscle force to stabilize digital joints, and are exposed to relatively lower joint contact stresses during stone tool use, due in part to an increase in the robusticity of metacarpals and phalanges in humans relative to chimpanzees. These analyses further suggest that Pan- or australopith-like pollical anatomy presents serious performance challenges to habitual tool use. Our data support the hypothesis that evolutionary increases in thumb length, robusticity, and thenar muscle mass enabled Homo to produce more force and to tolerate higher joint stresses during tool use.     

Variation in enamel thickness within the genus Homo

Recent humans and their fossil relatives are classified as having thick molar enamel, one of very few dental traits that distinguish hominins from living African apes. However, little is known about enamel thickness in the earliest members of the genus Homo, and recent studies of later Homo report considerable intra- and inter-specific variation. In order to assess taxonomic, geographic, and temporal trends in enamel thickness, we applied micro-computed tomographic imaging to 150 fossil Homo teeth spanning two million years. Early Homo postcanine teeth from Africa and Asia show highly variable average and relative enamel thickness (AET and RET) values. Three molars from South Africa exceed Homo AET and RET ranges, resembling the hyper thick Paranthropus condition. Most later Homo groups (archaic European and north African Homo, and fossil and recent Homo sapiens) possess absolutely and relatively thick enamel across the entire dentition. In contrast, Neanderthals show relatively thin enamel in their incisors, canines, premolars, and molars, although incisor AET values are similar to H. sapiens. Comparisons of recent and fossil H. sapiens reveal that dental size reduction has led to a disproportionate decrease in coronal dentine compared with enamel (although both are reduced), leading to relatively thicker enamel in recent humans. General characterizations of hominins as having ‘thick enamel’ thus oversimplify a surprisingly variable craniodental trait with limited taxonomic utility within a genus. Moreover, estimates of dental attrition rates employed in paleodemographic reconstruction may be biased when this variation is not considered. Additional research is necessary to reconstruct hominin dietary ecology since thick enamel is not a prerequisite for hard-object feeding, and it is present in most later Homo species despite advances in technology and food processing.     

New evidence for a 67,000-year-old human presence at Callao Cave, Luzon, Philippines

Documentation of early human migrations through Island Southeast Asia and Wallacea en route to Australia has always been problematic due to a lack of well-dated human skeletal remains. The best known modern humans are from Niah Cave in Borneo (40-42 ka), and from Tabon Cave on the island of Palawan, southwest Philippines (47 ± 11 ka). The discovery of Homo floresiensis on the island of Flores in eastern Indonesia has also highlighted the possibilities of identifying new hominin species on islands in the region. Here, we report the discovery of a human third metatarsal from Callao Cave in northern Luzon. Direct dating of the specimen using U-series ablation has provided a minimum age estimate of 66.7 ± 1 ka, making it the oldest known human fossil in the Philippines. Its morphological features, as well as size and shape characteristics, indicate that the Callao metatarsal definitely belongs to the genus Homo. Morphometric analysis of the Callao metatarsal indicates that it has a gracile structure, close to that observed in other small-bodied Homo sapiens. Interestingly, the Callao metatarsal also falls within the morphological and size ranges of Homo habilis and H. floresiensis. Identifying whether the metatarsal represents the earliest record of H. sapiens so far recorded anywhere east of Wallace’s Line requires further archaeological research, but its presence on the isolated island of Luzon over 65,000 years ago further demonstrates the abilities of humans to make open ocean crossings in the Late Pleistocene.