New hominin dental remains from the ∼2.04-1.95 Ma Drimolen Main Quarry, South Africa.

BACKGROUND: The Drimolen Palaeocave site is situated within the UNESCO Fossil Hominid Sites of South Africa World Heritage Area and has yielded numerous hominin fossils since its discovery in 1992. Most of these fossils are represented by isolated dental elements, which have been attributed to either of two distinct hominin genera, Paranthropus and Homo. AIM: This paper provides morphological descriptions for a further 19 specimens that have been recovered from the ∼2.04-1.95 Ma Drimolen Main Quarry (DMQ) deposits since 2008. This paper also discusses the two primary hypotheses used to explain Paranthropus robustus variation: sexual dimorphism, and micro-evolution within a lineage. SUBJECTS AND METHODS: These 19 fossils are represented by 47 dental elements and expand the sample of DMQ early Homo from 13 to 15, and the sample of Paranthropus robustus from 69 to 84. RESULTS: The evidence presented in this paper was found to be inconsistent with the sexual dimorphism hypothesis. CONCLUSION: Some support was found for the micro-evolution hypothesis.

The Ornaments of the Arma Veirana Early Mesolithic Infant Burial.

Personal ornaments are widely viewed as indicators of social identity and personhood. Ornaments are ubiquitous from the Late Pleistocene to the Holocene, but they are most often found as isolated objects within archaeological assemblages without direct evidence on how they were displayed. This article presents a detailed record of the ornaments found in direct association with an Early Mesolithic buried female infant discovered in 2017 at the site of Arma Veirana (Liguria, Italy). It uses microscopic, 3D, and positional analyses of the ornaments as well as a preliminary perforation experiment to document how they were perforated, used, and what led to their deposit as part of the infant's grave goods. This study provides important information on the use of beads in the Early Mesolithic, in general, as well as the relationship between beads and young subadults, in particular. The results of the study suggest that the beads were worn by members of the infant's community for a considerable period before they were sewn onto a sling, possibly used to keep the infant close to the parents while allowing their mobility, as seen in some modern forager groups. The baby was then likely buried in this sling to avoid reusing the beads that had failed to protect her or simply to create a lasting connection between the deceased infant and her community. Supplementary Information: The online version contains supplementary material available at 10.1007/s10816-022-09573-7.

Comment on "Origin of human bipedalism as an adaptation for locomotion on flexible branches".

Thorpe et al. (Reports, 1 June 2007, p. 1328) concluded that human bipedalism evolved from a type of bipedal posture they observed in extant orangutans with seemingly human-like extended knees. However, humans share knuckle-walking characters with African apes that are absent in orangutans. These are most parsimoniously explained by positing a knuckle-walking precursor to human bipedalism.

Molar enamel thickness in the chacma baboon, Papio ursinus (Kerr 1792).

Modern humans exhibit increasing relative enamel thickness from M1 to M3. Some biomechanical (basic lever) models predict that the more distal molars in humans encounter higher occlusal forces, and it has been postulated that this provides a functional explanation for the observed gradient in relative enamel thickness. However, constrained three-dimensional models and experimental observations suggest that there is a reduction in bite force potential from M1 to M3, which would be consistent with the tendency for humans to reduce the size of the distal molars. In this regard, it has been postulated that the distal increase in enamel thickness is a consequence of crown size reduction; thus, it is unnecessary to invoke functional scenarios to explain this phenomenon. We assess these competing proposals by examining relative enamel thickness in a catarrhine primate (Papio ursinus) that exhibits crown size increase from M1 to M3. The molar row of P. ursinus is positioned relatively far forward of the temporomandibular joint, which results in the baboon being able to exert relatively greater muscle forces during posterior biting in comparison to modern humans. Thus, a significant distalward gradient of increasing enamel thickness would be expected in P. ursinus according to the hypothesis that posits it to be functionally related to bite force. The present study reveals no significant difference in relative enamel thickness along the molar row in P. ursinus. This finding lends support to the notion that the relatively thicker enamel of human distal molars is related primarily to their reduction in size. This carries potential implications for the interpretation of enamel thickness in phylogenetic reconstructions: the relatively thick molar enamel shared by modern humans and some of our fossil relatives may not be strictly homologous, in that it may result from different underlying developmental mechanisms.

Integration, phylogeny, and the hominid cranial base.

Basicranial features were examined in catarrhine primates and early hominids in order to demonstrate how information about morphological integration can be incorporated into phylogenetic analysis. Hypotheses purporting to explain the functional and structural relationships of basicranial characters were tested using factor analysis. Characters found to be functionally or structurally related to each other were then further examined in order to determine whether there was evidence that they were phylogenetically independent. If phylogenetic independence could not be demonstrated, then the characters were presumed to be integrated and were grouped into a complex. That complex was then treated as if it were a single character for the purposes of cladistic analysis. Factor analysis revealed that five basicranial features may be structurally related to relative brain size in hominoids. Depending on how one defines phylogenetic independence, as few as two, or as many as all of those characters might be morphologically integrated. A cladistic analysis of early hominids based on basicranial features revealed that the use of integrated complexes had a substantial effect on the phylogenetic position of Australopithecus africanus, a species whose relationships are poorly resolved. Moreover, the use of complexes also had an effect on reanalyses of certain published cladistic data sets, implying that those studies might have been biased by patterns of basicranial integration. These results demonstrate that patterns of morphological integration need to be considered carefully in all morphology-based cladistic analyses, regardless of taxon or anatomical focus. However, an important caveat is that the functional and structural hypotheses tested here predicted much higher degrees of integration than were observed. This result warns strongly that hypotheses of integration must be tested before they can be adequately employed in phylogenetic analysis. The uncritical acceptance of an untested hypothesis of integration is likely to be as disruptive to a cladistic analysis as when integration is ignored.

Origin of human bipedalism: The knuckle-walking hypothesis revisited.

Some of the most long-standing questions in paleoanthropology concern how and why human bipedalism evolved. Over the last century, many hypotheses have been offered on the mode of locomotion from which bipedalism originated. Candidate ancestral adaptations include monkey-like arboreal or terrestrial quadrupedalism, gibbon- or orangutan-like (or other forms of) climbing and suspension, and knuckle-walking. This paper reviews the history of these hypotheses, outlines their predictions, and assesses them in light of current phylogenetic, comparative anatomical, and fossil evidence. The functional significance of characteristics of the shoulder and arm, elbow, wrist, and hand shared by African apes and humans, including their fossil relatives, most strongly supports the knuckle-walking hypothesis, which reconstructs the ancestor as being adapted to knuckle-walking and arboreal climbing. Future fossil discoveries, and a clear understanding of anthropoid locomotor anatomy, are required to ultimately test these hypotheses. If knuckle-walking was an important component of the behavioral repertoire of the prebipedal human ancestor, then we can reject scenarios on the origin of bipedalism that rely on a strictly arboreal ancestor. Moreover, paleoenvironmental data associated with the earliest hominins, and their close relatives, contradict hypotheses that place the agents of selection for bipedality in open savanna habitats. Existing hypotheses must explain why bipedalism would evolve from an ancestor that was already partly terrestrial. Many food acquisition and carrying hypotheses remain tenable in light of current evidence.

Evidence that humans evolved from a knuckle-walking ancestor.

Bipedalism has traditionally been regarded as the fundamental adaptation that sets hominids apart from other primates. Fossil evidence demonstrates that by 4.1 million years ago, and perhaps earlier, hominids exhibited adaptations to bipedal walking. At present, however, the fossil record offers little information about the origin of bipedalism, and despite nearly a century of research on existing fossils and comparative anatomy, there is still no consensus concerning the mode of locomotion that preceded bipedalism. Here we present evidence that fossils attributed to Australopithecus anamensis (KNM-ER 20419) and A. afarensis (AL 288-1) retain specialized wrist morphology associated with knuckle-walking. This distal radial morphology differs from that of later hominids and non-knuckle-walking anthropoid primates, suggesting that knuckle-walking is a derived feature of the African ape and human clade. This removes key morphological evidence for a Pan-Gorilla clade, and suggests that bipedal hominids evolved from a knuckle-walking ancestor that was already partly terrestrial.

The scaling of basicranial flexion and length.

Based on correlations between the cranial base angle (CBA) and the index of relative encephalization (IRE, calculated as the cubed root of brain volume divided by basicranial length), several recent studies have identified relative brain size as the factor most responsible for determining basicranial flexion in primates. IRE, however, scales with positive allometry relative to body mass, unlike the negatively allometric relationship between brain volume and body mass. This poses new questions concerning the factors underlying the correlation between IRE and CBA. Specifically, if basicranial flexion represents a spatial solution to the problem of housing a large brain within a neurocranium of limited size, then why is it that the problem is greatest in those species whose brains are smallest relative to body mass? To address this question, the scaling relationships of IRE and the measurements used to calculate it were examined in 87 primate species. It was found that the positive allometry of IRE is due to the fact that its denominator, basicranial length (BL), scales with very strong negative allometry relative to body mass. The scaling relationship of BL may reflect the fact that the noncortical components of the brain (i.e., diencephalon, mesencephalon, medulla) also scale with strong negative allometry relative to body mass, perhaps because of energetic constraints. Importantly, BL and these three brain components scale isometrically against each other. Thus, although cranial base flexion may be an adaptation to accommodate the size of the brain relative to basicranial length, the reason why that adaptation is necessary is not the evolution of a large brain, but rather the evolution of a short cranial base. In so far as basicranial length is affected by the strong negative allometry of the diencephalon, mesencephalon and medulla, the scaling relationships of these brain components are therefore indirectly responsible for the evolution of basicranial flexion.

Early hominid biogeography.

We examined the biogeographic patterns implied by early hominid phylogenies and compared them to the known dispersal patterns of Plio-Pleistocene African mammals. All recent published phylogenies require between four and seven hominid dispersal events between southern Africa, eastern Africa, and the Malawi Rift, a greater number of dispersals than has previously been supposed. Most hominid species dispersed at the same time and in the same direction as other African mammals. However, depending on the ages of critical hominid specimens, many phylogenies identify at least one hominid species that dispersed in the direction opposite that of contemporaneous mammals. This suggests that those hominids may have possessed adaptations that allowed them to depart from continental patterns of mammalian dispersal.

Kinematic data on primate head and neck posture: implications for the evolution of basicranial flexion and an evaluation of registration planes used in paleoanthropology.

Kinematic data on primate head and neck posture were collected by filming 29 primate species during locomotion. These were used to test whether head and neck posture are significant influences on basicranial flexion and whether the Frankfurt plane can legitimately be employed in paleoanthropological studies. Three kinematic measurements were recorded as angles relative to the gravity vector, the inclination of the orbital plane, the inclination of the neck, and the inclination of the Frankfurt plane. A fourth kinematic measurement was calculated as the angle between the neck and the orbital plane (the head-neck angle [HNA]). The functional relationships of basicranial flexion were examined by calculating the correlations and partial correlations between HNA and craniometric measurements representing basicranial flexion, orbital kyphosis, and relative brain size (Ross and Ravosa [1993] Am. J. Phys. Anthropol. 91:305-324). Significant partial correlations were observed between relative brain size and basicranial flexion and between HNA and orbital kyphosis. This indicates that brain size, rather than head and neck posture, is the primary influence on flexion, while the degree of orbital kyphosis may act to reorient the visual field in response to variation in head and neck posture. Regarding registration planes, the Frankfurt plane was found to be horizontal in humans but inclined in all nonhuman primates. In contrast, nearly all primates (including humans) oriented their orbits such that they faced anteriorly and slightly inferiorly. These results suggest that for certain functional craniometric studies, the orbital plane may be a more suitable registration plane than Frankfurt "Horizontal."

A reappraisal of early hominid phylogeny.

We report here on the results of a new cladistic analysis of early hominid relationships. Ingroup taxa included Australopithecus afarensis, Australopithecus africanus, Australopithecus aethiopicus, Australopithecus robustus, Australopithecus boisei, Homo habilis, Homo rudolfensis, Homo ergaster and Homo sapiens. Outgroup taxa included Pan troglodytes and Gorilla gorilla. Sixty craniodental characters were selected for analysis. These were drawn from the trait lists of other studies and our own observations. Eight parsimony analyses were performed that differed with respect to the number of characters examined and the manner in which the characters were treated. Seven employed ordered characters, and included analyses in which (1) taxa that were variable with respect to a character were coded as having an intermediate state, (2) characters with variable states in any taxon were excluded; (3) a variable taxon was coded as having the state exhibited by the majority of its hypodigm, (4) variable taxa were coded as missing data for that character, (5) some characters were considered irreversible, (6) masticatory characters were excluded, and (7) characters whose states were unknown in some taxa were excluded. In the final analysis, (8) all characters were unordered. All analyses were performed using PAUP 3.0s. Despite the fact that the eight analyses differed with respect to methodology, they produced several consistent results. All agreed that the "robust" australopithecines form a clade, A. afarensis is the sister taxon of all other hominids, and the genus Australopithecus, as conventionally defined, is paraphyletic. All eight also supported trees in which A. africanus is the sister taxon of a joint Homo+ "robust" clade, although in one analysis an equally parsimonious topology found A. africanus to be the sister of the "robust" species. In most analyses, the relationships of A. africanus and H. habilis were unstable, in the sense that their positions vary in trees that are marginally less parsimonious than the favored one. Trees in which "robust" australopithecines are paraphyletic were found to be extremely unparsimonious.