Paranthropus Research Articles

The DNH 7 endocast of Paranthropus robustus from Drimolen, South Africa: Reconsidering the functional significance of an enlarged occipital-marginal (O/M) sinus system in robust australopithecines.

This paper presents a detailed analysis of the endocast of one of the most complete Paranthropus robustus crania known, DNH 7, from the Drimolen site (South Africa), and compares it with the morphology of other australopithecine endocasts. We focus on endocranial volume, the impressions of cortical sulci, cranial sutures, and the pattern of cranial venous sinuses on the endocast. A noteworthy observation is the estimated endocranial capacity of 403 cm3, which is small for an adult Paranthropus. Fragmentary sulci identified in the frontal and temporal lobes of DNH 7 exhibit similarities with patterns observed in chimpanzees and gracile australopithecines. We observe the presence of a large remnant of an occipital-marginal sinus on DNH 7 and provide an updated table of 13 Paranthropus endocasts that are scorable for this trait, which reinforces the hypothesis that an enlarged occipital-marginal (O/M) sinus system was fixed across the three species of Paranthropus. In light of this, the possible functional significance of the occipital-marginal sinus system is reevaluated considering the ontogenetic development of cranial venous blood flow in human children. This leads us to hypothesize that the ontogenetic development of cranial blood flow in Paranthropus and Australopithecus africanus infants were different and to suggest that Taung 1 was the only A. africanus specimen known to have exhibited an enlarged O/M sinus system because it was an immature individual.

We the hunted

Classic depictions of human evolutionary ecology cast Homo as predator and other hominins, including Paranthropus robustus, as prey. Such hypotheses rest on a small number of fossils that exhibit evidence of carnivore predation, including the iconic SK 54 cranium from Swartkrans in South Africa. Here we demonstrate that the SK 54 cranium shares its closest affinities with H. erectus sensu lato rather than P. robustus. Demonstrating that Homo was prey for leopards at Swartkrans weakens the historically significant hypothesis that Homo was better able to avoid predation because of being behaviourally and technologically advanced compared to Paranthropus. Subsequent ideas about hominin palaeobiology derived from this hypothesis warrant reconsideration.

New fossil Bovidae (Mammalia: Artiodactyla) from Kromdraai Unit P, South Africa and their implication for biochronology and hominin palaeoecology

Kromdraai is a Plio-Pleistocene site located in the Cradle of Humankind (Gauteng Province, South Africa). It has produced diverse and abundant faunal assemblages and key hominin specimens like the holotype of Paranthropus robustus and early Homo. We provide the first taxonomic study of the Bovidae Family from the hominin-bearing Unit P at Kromdraai and discuss its potential to unravel its paleoecological context. We describe the presence of an unknown medium-sized buffalo (Syncerus sp.) that could be closely related to S. acoelotus. The bovid assemblage from Kromdraai Unit P combines older Plio-Pleistocene (Gazella gracilior, Makapania broomi, Numidocapra cf. porrocornutus) and younger Pleistocene taxa (Tragelaphus strepsiceros, Oreotragus oreotragus, Raphicerus campestris, Damaliscus lunatus). Overall, these bovid species indicate a grassland-dominated environment for Unit P. Comparisons with other Plio-Pleistocene South African sites indicate that Australopithecus was associated with woodland and closed-wet environment-adapted taxa, whereas Homo was found in association with bovid species adapted to open and dry environments. In contrast, the assemblages associated with Paranthropus show an extensive range of environmental adaptation among the bovids. The biochronology indicates that Kromdraai Unit P accumulated between 2.9 and 1.8 Ma. If Kromdraai Unit P were to be confirmed to date to the older end of this scale, it would provide the earliest appearance of Paranthropus robustus, Numidocapra, and Damaliscus in southern Africa.

Revisiting mandibular symphyseal shape in juvenile early hominins and modern humans using a deformation-based approach.

The juvenile mandible is important in the investigation of ontogenetic and evolutionary changes among early hominins. We revisit the mandibular symphysis in juvenile specimens of Australopithecus africanus and Paranthropus robustus with two main contributions. First, we employ, for the first time, methods of computational anatomy to model complex symphyseal shape differences. Second, we present new fossil evidence from Kromdraai to improve our knowledge of symphyseal morphology. We describe differences between shapes by landmark-free diffeomorphism needed to align them. We assess which features of the mandibular symphysis best discriminate the juvenile symphysis in these fossil species, relative to the intraspecific variation observed among modern humans. Our approach eliminates potential methodological inconsistencies with traditional approaches (i.e., the need for homologous anatomical landmarks, assumption of linearity). By enabling detailed comparisons of complex shapes in juvenile mandibles, our proposed approach offers new perspectives for more detailed comparisons among Australopithecus, Paranthropus and early Homo.

Fossil micromammals and the palaeoenvironments of the Paranthropus robustus site of Cooper’s Cave

ABSTRACT The Cradle of Humankind (Gauteng, South Africa) provides an important fossil record of the evolutionary history of PlioPleistocene hominins. Cooper’s Cave deposits have yielded a rich fossil faunal assemblage, as well as six remains attributed to Paranthropus robustus. This study provides the first taxonomic, taphonomic and palaeoecological description of the micromammal material from the 1.4 Ma assemblage of Cooper’s D. The taphonomic signature of the assemblage indicates an accumulation by tytonid owls (probably Tyto alba) and advanced postdepositional disturbance probably related to trampling by the occupants of the cave, sorting of the bones along slope, and burying. The taxonomic analysis undertaken here at genus level describes at least 22 taxa of small mammals, including one extinct genus Proodontomys. This assemblage is dominated by Mystromys and Otomys, two rodent genera adapted to grassland habitats which are among the most common among pliopleistocene micromammal faunas from the region. The palaeoecological analysis suggests an open landscape with a predominance of grassland and savanna vegetation, and the proximity of rocky outcrops and a perennial river. These results support previous indications of a shift in the African climate and vegetation towards more open habitats during the Early Pleistocene.

Estimates of absolute crown strength and bite force in the lower postcanine dentition of Gigantopithecus blacki

Gigantopithecus blacki is hypothesized to have been capable of processing mechanically challenging foods, which likely required this species to have high dental resistance to fracture and/or large bite force. To test this hypothesis, we used two recently developed approaches to estimate absolute crown strength and bite force of the lower postcanine dentition. Sixteen Gigantopithecus mandibular permanent cheek teeth were scanned by micro-computed tomography. From virtual mesial cross-sections, we measured average enamel thickness and bi-cervical diameter to estimate absolute crown strength, and cuspal enamel thickness and dentine horn angle to estimate bite force. We compared G. blacki with a sample of extant great apes (Pan, Pongo, and Gorilla) and australopiths (Australopithecus anamensis, Australopithecus afarensis, Australopithecus africanus, Paranthropus robustus, and Paranthropus boisei). We also evaluated statistical differences in absolute crown strength and bite force between the premolars and molars for G. blacki. Results reveal that molar crown strength is absolutely greater, and molar bite force absolutely higher, in G. blacki than all other taxa except P. boisei, suggesting that G. blacki molars have exceptionally high resistance to fracture and the ability to generate exceptionally high bite force. In addition, G. blacki premolars have comparable absolute crown strength and larger bite force capabilities compared with its molars, implying possible functional specializations in premolars. The dental specialization of G. blacki could thus represent an adaptation to further facilitate the processing of mechanically challenging foods. While it is currently not possible to determine which types of foods were actually consumed by G. blacki through this study, direct evidence (e.g. dental chipping and microwear) left by the foods eaten by G. blacki could potentially lead to greater insights into its dietary ecology.

An updated analysis of hominin phylogeny with an emphasis on re-evaluating the phylogenetic relationships of Australopithecus sediba

The discovery and description of Australopithecus sediba has reignited the debate over the evolutionary history of the australopiths and the genus Homo. It has been suggested that A. sediba may be an ancestor of Homo because it possesses a mosaic of derived Homo-like and primitive australopith-like traits. However, an alternative hypothesis proposes that the majority of the purported Homo-like craniodental characters can be attributed to the juvenile status of the type specimen, MH1. We conducted an independent character assessment of the craniodental morphology of A. sediba, with particular emphasis on evaluating whether the ontogenetic status of MH1 may have affected its purported Homo-like characteristics. In doing so, we have also expanded fossil hypodigms to incorporate the new Australopithecus anamensis cranium from Woranso-Mille (MRD-VP-1/1), as well as recently described Paranthropus robustus cranial remains from Drimolen (DNH 7, DNH 155). Morphological character data were analyzed using both standard parsimony and Bayesian techniques. In addition, we conducted a series of Bayesian analyses constrained to evaluate the hypothesis that Australopithecus africanus and A. sediba are sister taxa. Based on the results of the parsimony and Bayesian analyses, we could not reject the hypothesis that A. sediba shares its closest phylogenetic affinities with the genus Homo. Therefore, based on currently available craniodental evidence, we conclude that A. sediba is plausibly the terminal end of a lineage that shared a common ancestor with the earliest representatives of Homo. We caution, however, that the discovery of new A. sediba fossils preserving adult cranial morphology or the inclusion of postcranial characters may ultimately necessitate a re-evaluation of this hypothesis.

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.

Morphology of early hominid hand in context of tool making

This article will focus on morphology of hand early hominids. We already have not many hands fossils, but this evidences can help us to interpret early hominid`s potential of tools making. We will talk about Australopithecus afarensis, Australopithecus africanus, Australopithecus sediba, Paranthropus robustus and Homo habilis. Early hominid research in Africa began in earnest in the second part of the 20th century, decades after the discovery of the Taung Child by Raymond Dart in 1924. After Dart’s momentous discovery finally gained widespread acceptance, scientists at last came to understand that human ancestry began in Africa rather than the Far East as was previously believed. Unfortunately there are very few carpal bones dating to this formative period of paleoanthropological research. Real interest in the development and implications of hominid hand morphology emerged only after the discovery of stone stools at Lomekwi 3 by Sonia Harmand in the 2010s. The discovery of 3 million year old tools fundamentally altered our understanding of the timeline of human cultural evolution. In addition to raising a number of pressing questions about the origins and definition of the genus Homo, Lomekwi 3 also radically reworked our understanding of the kinds of hand morphologies required for the production of stone tools. This brings us to the central question of this paper, namely, «which morphological features in early hominid hands are potentially indicative of tool making?». When we are talking about Lomekwi its very important to understand that the stone tools found at the site have not yet been associated with any paleoanthropological remains. Nonetheless, the 3.3 million year old date of the Lomekwi tools calls into question whether or not tool making can be said to be a trait unique to the genus Homo. We will see that by interpreting the carpal bones of the various early hominids present 3.3. million years ago we are forced not only to abandon outmoded assumptions about uniquely Homo traits, but to also reconsider the cognitive and mechanical capabilities of previously underestimated early hominid groups. Our goal is to determine which groups of early hominids may have been capable of producing the Oldowan and Lomekwi toolkits on the basis of functional morphology alone. Because we don’t find hominid remains holding tools, we need to make educated guesses about who made them. We can look at hand morphology, and its development, to attribute early tools to the particular hominids who may have made them. Key words: archaeology, paleoanthropology, evolution, Australopithecus, Paranthropus, Early hominids.

The Kromdraai early hominin-bearing site. A review of recent findings

The Kromdraai archaeological site is located in a fossiliferous paleokarst situated in the UNESCO World Heritage Site referred to as the “Cradle of Humankind” in the Gauteng Province of South Africa. Kromdraai is noteworthy because it features among the three southern African early hominin-bearing sites considered to represent distinct temporal periods within the same stratigraphic succession. Kromdraai also yielded a partial skull and dentition (TM 1517) in 1938 that was designated as the holotype of a new genus and species, Paranthropus robustus. Although the hominin fossil assemblage collected from Kromdraai between 1938 and 2014 is rather paltry, morphometric and cladistic analyses of this material suggested that it represented a somewhat less-derived form of P. robustus than the considerably larger assemblage from the nearby site of Swartkrans. However, the geochronological and biotic relationships among the P. robustus-bearing sites in South Africa are not resolved. Since 2014, the previously unknown, albeit densely fossiliferous Unit P produced 51 individually catalogued hominin fossils (36 craniodental and 15 postcranial) that currently represent 13% of the faunal assemblage from this unit with a minimum number of 10 juvenile and 9 adult individuals. P. robustus and early Homo coexisted at the time of the accumulation of Unit P at Kromdraai, with a relative abundance of 89% and 11%, respectively. P. robustus and early Homo are associated with a highly diverse fauna that includes several water-dependent species, and a large variety of bovid and carnivore taxa. Biochronological data from Unit P and an interval of reversed polarity measured in younger deposits above it are interpreted in the context of the regional chronological framework to infer that it represents a deposit that was likely accumulated prior to 2 Ma.

New fossils from Kromdraai and Drimolen, South Africa, and their distinctiveness among Paranthropus robustus

Most fossil hominin species are sampled with spatial, temporal or anatomical biases that can hinder assessments of their paleodiversity, and may not yield genuine evolutionary signals. We use new fossils from the Kromdraai (Unit P) and Drimolen sites (South Africa) to provide insights into the paleodiversity of the Lower Pleistocene robust australopith, Paranthropus robustus. Our focus is the morphology of the temporal bone and the relationships between size and shape (allometry) of the semi-circular canals (SCC), an aspect that has not yet been investigated among southern African australopiths. We find significant size and shape SCC differences between P. robustus from Kromdraai, Drimolen and Swartkrans. This site-related variation is consistent with other differences observed on the temporal bone. P. robustus from Kromdraai Unit P is distinctive because of its smaller temporal bone and SCC, and its proportionally less developed posterior SCC, independently of age and sex. We emphasize the importance of allometry to interpret paleodiversity in P. robustus as either the consequence of differences in body size, or as yet unknown factors. Some features of the inner ear of P. robustus represent directional selection soon after its origin, whereas the size and shape variations described here may result from evolutionary changes.

New hominin dental remains from the Drimolen Main Quarry, South Africa (1999–2008)

AbstractObjectivesTwenty‐four dental specimens from the Drimolen Main Quarry (DMQ) are described. This increases the number of DMQ Paranthropus robustus specimens from 48 to 63 and DMQ Homo specimens from 8 to 12. This allows reassessment of the proposed differences between the DMQ P. robustus assemblage and that of Swartkrans. Analysis conducted assesses intraspecific and inter‐locality variation.Materials and MethodsWe examined the P. robustus and early Homo assemblages from South Africa. Morphology was observed using a hand lens and a binocular microscope. Mesiodistal and buccolingual measurements were taken using plastic‐tipped calipers. Summary statistics were generated and patterns of variability in P. robustus were assessed through box plots and Mann–Whitney U tests.ResultsComparison between the expanded DMQ and Swartkrans P. robustus assemblages demonstrates overlap in size. Ten dental variables show statistically significant differences.DiscussionThe expanded P. robustus sample allowed us to re‐examine previous analyses of differences in tooth size between the samples. While analyses presented here show a high degree of overlap in the MD and BL dimensions of the two assemblages, significant differences were found in the mean values of these variables in the postcanine maxillary teeth—consistent with previous analyses. Two current hypotheses may explain this pattern: 1) dental size increase through the P. robustus lineage or 2) different sample composition between the two sites. Small sample sizes for all permanent dental classes in the DMQ assemblage represents a limitation on this analysis and interpretations thereof. Any addition to the DMQ or the Swartkrans samples may alter these results.

Fossil Bovidae from the Hominini-bearing site of Cooper’s D (Bloubank Valley, South Africa): implications for Paranthropus robustus Broom, 1938 and early Homo Linnaeus, 1758 habitat preferences

The Cooper’s D deposit has been dated by U-Pb at a maximum age of 1.37 Ma ± 0.113 and has yielded seven Hominini remains, six of them are attributed to the Southern African endemic species Paranthropus robustus Broom, 1938. However, the taxonomic composition of the faunal assemblage recovered at this site, especially ruminants, remains poorly understood. This paper updates the previous palaeontological and biochronological works and provides the first taxonomic description and paleoecological analysis of the whole bovid material from Cooper’s D. We propose a minimum age of 1.0 Ma for the Cooper’s D bone accumulation. Thus, we conclude that the assemblage accumulated between 1.375 ± 0.113 and 1.0 Ma based on the complementary results obtained from radiometric dates and biochronology. The paleoecological analysis suggests that Cooper’s D is likewise dominated by grassland with sparse covered habitat and probable water sources in the vicinity. Finally, we show a strong statistically significant negative correlation between the presence of grassland-adapted bovid species and P. robustus. This result indicates that P. robustus was more likely to be an eurytopic species and dietary generalist than an exclusive grazer.

Calcar femorale variation in extant and fossil hominids: Implications for identifying bipedal locomotion in fossil hominins

The calcar femorale is an internal bony structure of the proximal femur considered to be functionally related to bipedal locomotion. Among extant primates, the presence of a calcar femorale has been so far documented in extant humans and Pan and, among extinct hominins, in the Late Miocene Orrorin, in a Pliocene Australopithecus, and in a Middle Pleistocene Homo specimen. Using high-resolution microcomputed tomography, we investigated the occurrence and morphology (i.e., shape, location, and size) of the calcar femorale in an adult sample of extant humans, Pan troglodytes, Gorilla gorilla, Pongo sp., and Papio ursinus. We also investigated for the first time the occurrence and morphology of a calcar femorale in the adult proximal femoral remains of a Late Miocene great ape (Rudapithecus) and five Plio-Pleistocene hominins from Southern and Eastern Africa (Australopithecus and Paranthropus). We took four measurements: periosteal-to-tip maximum length, maximum length excluding cortical thickness, maximum vertical height, and the distance between the most anterior and posterior limits of the root. To allow for intergeneric comparisons, estimated body size was used to standardize all measurements. Nine of 10 extant humans have a well-developed calcar femorale. Among the African apes, 6 of 10 Pan and 6 of 10 Gorilla also show a distinct calcar femorale. In Pongo (n = 9), it is only present in one captive individual. None of the five investigated Papio specimens show any trace of this structure. Only calcar femorale height, which is systematically taller and extends into the lower part of the lesser trochanter, discriminates humans from extant great apes, except for one Gorilla. The calcar femorale was absent in one Paranthropus robustus and variably developed in all other investigated fossils. These results indicate that this structure cannot be considered as a diagnostic feature of habitual bipedal locomotion and emphasize the need for further investigations of its functional role.

A new early hominin calcaneus from Kromdraai (South Africa).

The Kromdraai site in South Africa has yielded numerous early hominin fossils since 1938. As a part of recent excavations within Unit P, a largely complete early hominin calcaneus (KW 6302) was discovered. Due to its role in locomotion, the calcaneus has the potential to reveal important form/function relationships. Here, we describe KW 6302 and analyze its preserved morphology relative to human and nonhuman ape calcanei, as well as calcanei attributed to Australopithecus afarensis, Australopithecus africanus, Australopithecus sediba, Homo naledi, and the Omo calcaneus (either Paranthropus or early Homo). KW 6302 calcaneal morphology is assessed using numerous quantitative metrics including linear measures, calcaneal robusticity index, relative lateral plantar process position, Achilles tendon length reconstruction, and a three-dimensional geometric morphometric sliding semilandmark analysis. KW 6302 exhibits an overall calcaneal morphology that is intermediate between humans and nonhuman apes, although closer to modern humans. KW 6302 possesses many traits that indicate it was likely well-adapted for terrestrial bipedal locomotion, including a relatively flat posterior talar facet and a large lateral plantar process that is similarly positioned to modern humans. It also retains traits that indicate that climbing may have remained a part of its locomotor repertoire, such as a relatively gracile tuber and a large peroneal trochlea. Specimens from Kromdraai have been attributed to either Paranthropus robustus or early Homo; however, there are no definitively attributed calcanei for either genus, making it difficult to taxonomically assign this specimen. KW 6302 and the Omo calcaneus, however, fall outside the range of expected variation for an extant genus, indicating that if the Omo calcaneus was Paranthropus, then KW 6302 would likely be attributed to early Homo (or vice versa).

Hominin Femur finding from Banjarejo: Its morphological character and taxonomical position

Recently, Banjarejo became an important prehistoric site for Quaternary research because of its significant faunal and hominin remains. This study aims to describe a new finding of the hominin femur specimen from the site and its taxonomical position in human evolution. The specimen was identified by morphological and metric descriptions of the external feature of the femur. Then, comparative study to Homo erectus (n=2), Homo neanderthalensis (n=8), Homo heidelbergensis (n=1), prehistoric Homo sapiens (n=44), Australopithecus africanus (n=1), Paranthropus robustus (n=2), also non-human primate including Pongo (n=1), Macaca (n=1) and Gibbon (n=1) using bivariate and multivariate statistical analysis presented the specimen in the evolutionary perspective. This study shows the morphological and metric character of Banjarejo specimen located between Homo erectus and prehistoric Homo sapiens population sample. Further study should be addressed to investigate the cultural and chronological context of the hominin specimen.

Cochlear shape distinguishes southern African early hominin taxa with unique auditory ecologies

Insights into potential differences among the bony labyrinths of Plio-Pleistocene hominins may inform their evolutionary histories and sensory ecologies. We use four recently-discovered bony labyrinths from the site of Kromdraai to significantly expand the sample for Paranthropus robustus. Diffeomorphometry, which provides detailed information about cochlear shape, reveals size-independent differences in cochlear shape between P. robustus and Australopithecus africanus that exceed those among modern humans and the African apes. The cochlea of P. robustus is distinctive and relatively invariant, whereas cochlear shape in A. africanus is more variable, resembles that of early Homo, and shows a degree of morphological polymorphism comparable to that evinced by modern species. The curvature of the P. robustus cochlea is uniquely derived and is consistent with enhanced sensitivity to low-frequency sounds. Combined with evidence for selection, our findings suggest that sound perception shaped distinct ecological adaptations among southern African early hominins.

New evidence of bone tool use by Early Pleistocene hominins from Cooper’s D, Bloubank Valley, South Africa

Bone tool-use by Early Pleistocene hominins is at the centre of debates in human evolution. It is especially the case in South Africa, where 102 bone tools have been described from four Early Stone Age archaeological sites, which have yielded Oldowan and possibly Acheulean artefacts, as well as Paranthropus robustus and early Homo remains. Here we describe a bone tool from Cooper’s D. The deposit, dated between 1.4 and 1.0 Ma, has yielded seven P. robustus remains and 50 stone artefacts. Our results highlight similarities in morphology and use-wear patterns between the Cooper’s D bone tool and those previously identified at nearby Sterkfontein, Swartkrans, Kromdraai and Drimolen. Our findings increase the number of Early Stone Age bone tools and provide further evidence of their association with P. robustus. They suggest P. robustus had the cognitive capacities to develop this cultural adaptation and the manipulative abilities to implement it.

Paranthropus robustus tooth chipping patterns do not support regular hard food mastication

Paranthropus robustus tooth chipping patterns do not support regular hard food mastication

Relative tooth size, Bayesian inference, and Homo naledi.

Size-corrected tooth crown measurements were used to estimate phenetic affinities among Homo naledi (~335-236 ka) and 11 other Plio-Pleistocene and recent species. To assess further their efficacy, and identify dental evolutionary trends, the data were then quantitatively coded for phylogenetic analyses. Results from both methods contribute additional characterization of H. naledi relative to other hominins. After division by their geometric mean, scaled mesiodistal and buccolingual dimensions were used in tooth size apportionment analysis to compare H. naledi with Australopithecus africanus, A. afarensis, Paranthropus robustus, P. boisei, H. habilis, H. ergaster, H. erectus, H. heidelbergensis, H. neanderthalensis, H. sapiens, and Pan troglodytes. These data produce equivalently scaled samples unaffected by interspecific size differences. The data were then gap-weighted for Bayesian inference. Congruence in interspecific relationships is evident between methods, and with many inferred from earlier systematic studies. However, the present results place H. naledi as a sister taxon to H. habilis, based on a symplesiomorphic pattern of relative tooth size. In the preferred Bayesian phylogram, H. naledi is nested within a clade comprising all Homo species, but it shares some characteristics with australopiths and, particularly, early Homo. Phylogenetic analyses of relative tooth size yield information about evolutionary dental trends not previously reported in H. naledi and the other hominins. Moreover, with an appropriate model these data recovered plausible evolutionary relationships. Together, the findings support recent study suggesting H. naledi originated long before the geological date of the Dinaledi Chamber, from which the specimens under study were recovered.