'Snakes and ladders' in paleoanthropology: From cognitive surprise to skillfulness a million years ago.

A paradigmatic account may suffice to explain behavioral evolution in early Homo. We propose a parsimonious account that (1) could explain a particular, frequently-encountered, archeological outcome of behavior in early Homo - namely, the fashioning of a Paleolithic stone 'handaxe' - from a biological theoretic perspective informed by the free energy principle (FEP); and that (2) regards instances of the outcome as postdictive or retrodictive, circumstantial corroboration. Our proposal considers humankind evolving as a self-organizing biological ecosystem at a geological time-scale. We offer a narrative treatment of this self-organization in terms of the FEP. Specifically, we indicate how 'cognitive surprises' could underwrite an evolving propensity in early Homo to express sporadic unorthodox or anomalous behavior. This co-evolutionary propensity has left us a legacy of Paleolithic artifacts that is reminiscent of a 'snakes and ladders' board game of appearances, disappearances, and reappearances of particular archeological traces of Paleolithic behavior. When detected in the Early and Middle Pleistocene record, anthropologists and archeologists often imagine evidence of unusual or novel behavior in terms of early humankind ascending the rungs of a figurative phylogenetic 'ladder' - as if these corresponded to progressive evolution of cognitive abilities that enabled incremental achievements of increasingly innovative technical prowess, culminating in the cognitive ascendancy of Homo sapiens. The conjecture overlooks a plausible likelihood that behavior by an individual who was atypical among her conspecifics could have been disregarded in a community of Hominina (for definition see Appendix 1) that failed to recognize, imagine, or articulate potential advantages of adopting hitherto unorthodox behavior. Such failure, as well as diverse fortuitous demographic accidents, would cause exceptional personal behavior to be ignored and hence unremembered. It could disappear by a pitfall, down a 'snake', as it were, in the figurative evolutionary board game; thereby causing a discontinuity in the evolution of human behavior that presents like an evolutionary puzzle. The puzzle discomforts some paleoanthropologists trained in the natural and life sciences. They often dismiss it, explaining it away with such self-justifying conjectures as that, maybe, separate paleospecies of Homo differentially possessed different cognitive abilities, which, supposedly, could account for the presence or absence in the Pleistocene archeological record of traces of this or that behavioral outcome or skill. We argue that an alternative perspective - that inherits from the FEP and an individual's 'active inference' about its surroundings and of its own responses - affords a prosaic, deflationary, and parsimonious way to account for appearances, disappearances, and reappearances of particular behavioral outcomes and skills of early humankind.

The driving force behind tool-stone selection in the African Middle Stone Age.

In the Stone Age, the collection of specific rocks was the first step in tool making. Very little is known about the choices made during tool-stone acquisition. Were choices governed by the knowledge of, and need for, specific properties of stones? Or were the collected raw materials a mere by-product of the way people moved through the landscape? We investigate these questions in the Middle Stone Age (MSA) of southern Africa, analyzing the mechanical properties of tool-stones used at the site Diepkloof Rock Shelter. To understand knapping quality, we measure flaking predictability and introduce a physical model that allows calculating the relative force necessary to produce flakes from different rocks. To evaluate their quality as finished tools, we investigate their resistance during repeated use activities (scraping or cutting) and their strength during projectile impacts. Our findings explain tool-stone selection in two emblematic periods of the MSA, the Still Bay and Howiesons Poort, as being the result of a deep understanding of these mechanical properties. In both cases, people chose those rocks, among many others, that allowed the most advantageous trade-off between anticipated properties of finished tools and the ease of acquiring rocks and producing tools. The implications are an understanding of African MSA toolmakers as engineers who carefully weighed their choices taking into account workability and the quality of the tools they made.

The brain of Homo habilis: Three decades of paleoneurology.

In 1987, Phillip Tobias published a comprehensive anatomical analysis of the endocasts attributed to Homo habilis, discussing issues dealing with brain size, sulcal patterns, and vascular traces. He suggested that the neuroanatomy of this species evidenced a clear change toward many cerebral traits associated with our genus, mostly when concerning the morphology of the frontal and parietal cortex. After more than 30 years, the fossil record associated with this taxon has not grown that much, but we have much more information on cranial and brain biology, and we are using a larger array of digital methods to investigate the paleoneurological variation observed in the human genus. Brain volume, the size of the frontal lobe, or the gross hemispheric asymmetries are still relevant issues, but they are considered to be less central than before. More attention is instead being paid to the cortical organization, the relationships with the cranial architecture, and the influence of molecular or ecological factors. Although the field of paleoneurology can currently count on a larger range of tools and principles, there is still a general lack of anatomical information on many endocranial traits. This aspect is probably crucial for the agenda of paleoneurology. More importantly, the whole science is undergoing a delicate change, because of the growing influence of the social environment. In this sense, the disciplines working with fossils (and, in particular, with brain evolution) should take particular care to maintain a healthy professional situation, avoiding an excess of speculation and overstatement.

Archaeological adhesives made from Podocarpus document innovative potential in the African Middle Stone Age.

Studying the earliest archaeological adhesives has implications for our understanding of human cognition. In southern Africa, the oldest adhesives were made by Homo sapiens in the Middle Stone Age. Chemical studies have shown that these adhesives were made from a local conifer of the Podocarpaceae family. However, Podocarpus does not exude resin, nor any other substance that could have been recognized as having adhesive properties. Therefore, it remains unknown how these adhesives were made. This study investigates how Podocarpus adhesives can be made, comparing their mechanical properties with other naturally available adhesives. We found that Podocarpus tar can only be made by dry distillation of leaves, requiring innovative potential, skill, and knowledge. This contrasts with our finding that the Middle Stone Age environment was rich in substances that can be used as adhesives without such transformation. The apparent preference for Podocarpus tar may be explained by its mechanical properties. We found it to be superior to all other substances in terms of its adhesive properties. In addition, the condensation method that allows producing it can be recognized accidentally, as the processes take place above ground and can be triggered accidentally. Our findings have implications for establishing a link between technology and cognition in the Middle Stone Age.

Dental data challenge the ubiquitous presence of Homo in the Cradle of Humankind.

The origins of Homo, as well as the diversity and biogeographic distribution of early Homo species, remain critical outstanding issues in paleoanthropology. Debates about the recognition of early Homo, first appearance dates, and taxonomic diversity within Homo are particularly important for determining the role that southern African taxa may have played in the origins of the genus. The correct identification of Homo remains also has implications for reconstructing phylogenetic relationships between species of Australopithecus and Paranthropus, and the links between early Homo species and Homo erectus. We use microcomputed tomography and landmark-free deformation-based three-dimensional geometric morphometrics to extract taxonomically informative data from the internal structure of postcanine teeth attributed to Early Pleistocene Homo in the southern African hominin-bearing sites of Sterkfontein, Swartkrans, Drimolen, and Kromdraai B. Our results indicate that, from our sample of 23 specimens, only 4 are unambiguously attributed to Homo, 3 of them coming from Swartkrans member 1 (SK 27, SK 847, and SKX 21204) and 1 from Sterkfontein (Sts 9). Three other specimens from Sterkfontein (StW 80 and 81, SE 1508, and StW 669) approximate the Homo condition in terms of overall enamel-dentine junction shape, but retain Australopithecus-like dental traits, and their generic status remains unclear. The other specimens, including SK 15, present a dominant australopith dental signature. In light of these results, previous dietary and ecological interpretations can be reevaluated, showing that the geochemical signal of one tooth from Kromdraai (KB 5223) and two from Swartkrans (SK 96 and SKX 268) is consistent with that of australopiths.

An ancient cranium from Dmanisi: Evidence for interpersonal violence, disease, and possible predation by carnivores on Early Pleistocene Homo.

Five well-preserved individuals from Dmanisi represent a paleodeme attributed to early Homo. Here we provide a case study of the D2280 adult cranium, which presents four oval-shaped lesions on the frontal, left parietal, and occipital bones. Several conditions are considered as possibly contributing to this pathology, including trauma, cysts, metastatic cancer, and infectious disease. One large but shallow depression on the left parietal bone has slightly elevated boundaries. Imaging reveals inner and outer tables that are reciprocally concave, so that the diploë is diminished or completely absent. This lesion is very likely a result of traumatic injury. Two additional depressions on the left side frontal and occipital bones may also be attributed to blunt force trauma. Such injuries stem from a variety of causes, but interpersonal violence may well be implicated. Based on the location and structure of a fourth lesion on the right-side frontal bone, we advance a possible diagnosis of treponemal disease. Lesions on the cranium and specifically on the frontal bone are common in treponemal disease. The condition develops as a periostitis, which eventually results in the destruction of the osteoperiosteal border of the cranial outer table and rarely involves the inner table. Additional perforations on the occipital bone are interpreted as perimortem damage resulting from predation by carnivores or scavenging by birds. If our reading of this cranial evidence can be confirmed, then D2280 documents one of the earliest instances of blunt force trauma in the Homo lineage. Dmanisi may also reveal the presence of treponemal disease in a population dated ca. 1.77 Ma ago. These findings bear on the social behavior of ancient humans and also the impact of infectious diseases on their survival.

A new absolute date from Swartkrans Cave for the oldest occurrences of Paranthropus robustus and Oldowan stone tools in South Africa.

The Early Pleistocene site of Swartkrans in South Africa's Cradle of Humankind World Heritage Site has been significant for our understanding of the evolution of both early Homo and Paranthropus, as well as the earliest archaeology of southern Africa. Previous attempts to improve a faunal age estimate of the earliest deposit, Member 1, had produced results obtained with uranium-lead dating (U-Pb) on flowstones and cosmogenic burial dating of quartz, which placed the entire member in the range of >1.7/1.8 Ma and <2.3 Ma. In 2014, two simple burial dates for the Lower Bank, the earliest unit within Member 1, narrowed its age to between ca. 1.8 Ma and 2.2 Ma. A new dating program using the isochron method for burial dating has established an absolute age of 2.22 ± 0.09 Ma for a large portion of the Lower Bank, which can now be identified as containing the earliest Oldowan stone tools and fossils of Paranthropus robustus in South Africa. This date agrees within one sigma with the U-Pb age of 2.25 ± 0.08 Ma previously published for the flowstone underlying the Lower Bank and confirms a relatively rapid rate of accumulation for a large portion of the talus.

The Middle to Later Stone Age transition at Panga ya Saidi, in the tropical coastal forest of eastern Africa.

The Middle to Later Stone Age transition is a critical period of human behavioral change that has been variously argued to pertain to the emergence of modern cognition, substantial population growth, and major dispersals of Homo sapiens within and beyond Africa. However, there is little consensus about when the transition occurred, the geographic patterning of its emergence, or even how it is manifested in the stone tool technology that is used to define it. Here, we examine a long sequence of lithic technological change at the cave site of Panga ya Saidi, Kenya, that spans the Middle and Later Stone Age and includes human occupations in each of the last five Marine Isotope Stages. In addition to the stone artifact technology, Panga ya Saidi preserves osseous and shell artifacts, enabling broader considerations of the covariation between different spheres of material culture. Several environmental proxies contextualize the artifactual record of human behavior at Panga ya Saidi. We compare technological change between the Middle and Later Stone Age with on-site paleoenvironmental manifestations of wider climatic fluctuations in the Late Pleistocene. The principal distinguishing feature of Middle from Later Stone Age technology at Panga ya Saidi is the preference for fine-grained stone, coupled with the creation of small flakes (miniaturization). Our review of the Middle to Later Stone Age transition elsewhere in eastern Africa and across the continent suggests that this broader distinction between the two periods is in fact widespread. We suggest that the Later Stone Age represents new short use-life and multicomponent ways of using stone tools, in which edge sharpness was prioritized over durability.

Biomechanics of the human thumb and the evolution of dexterity.

Systematic tool production and use is one of humanity's defining characteristics, possibly originating as early as >3 million years ago.1-3 Although heightened manual dexterity is considered to be intrinsically intertwined with tool use and manufacture, and critical for human evolution, its role in the emergence of early culture remains unclear. Most previous research on this question exclusively relied on direct morphological comparisons between early hominin and modern human skeletal elements, assuming that the degree of a species' dexterity depends on its similarity with the modern human form. Here, we develop a new approach to investigate the efficiency of thumb opposition, a fundamental component of manual dexterity, in several species of fossil hominins. Our work for the first time takes into account soft tissue as well as bone anatomy, integrating virtual modeling of musculus opponens pollicis and its interaction with three-dimensional bone shape form. Results indicate that a fundamental aspect of efficient thumb opposition appeared approximately 2 million years ago, possibly associated with our own genus Homo, and did not characterize Australopithecus, the earliest proposed stone tool maker. This was true also of the late Australopithecus species, Australopithecus sediba, previously found to exhibit human-like thumb proportions. In contrast, later Homo species, including the small-brained Homo naledi, show high levels of thumb opposition dexterity, highlighting the increasing importance of cultural processes and manual dexterity in later human evolution.

A genotype:phenotype approach to testing taxonomic hypotheses in hominids.

Paleontology has long relied on assumptions about the genetic and developmental influences on skeletal variation. The last few decades of developmental genetics have elucidated the genetic pathways involved in making teeth and patterning the dentition. Quantitative genetic analyses have refined this genotype:phenotype map even more, especially for primates. We now have the ability to define dental traits with a fair degree of fidelity to the underlying genetic architecture; for example, the molar module component (MMC) and the premolar-molar module (PMM) that have been defined through quantitative genetic analyses. We leverage an extensive dataset of extant and extinct hominoid dental variation to explore how these two genetically patterned phenotypes have evolved through time. We assess MMC and PMM to test the hypothesis that these two traits reveal a more biologically informed taxonomy at the genus and species levels than do more traditional measurements. Our results indicate that MMC values for hominids fall into two categories and that Homo is derived compared with earlier taxa. We find a more variable, species-level pattern for PMM. These results, in combination with previous research, demonstrate that MMC reflects the phenotypic output of a more evolutionarily stable, or phylogenetically congruent, genetic mechanism, and PMM is a reflection of a more evolutionarily labile mechanism. These results suggest that the human lineage since the split with chimpanzees may not represent as much genus-level variation as has been inferred from traits whose etiologies are not understood.

The upper limb of Paranthropus boisei from Ileret, Kenya.

Paranthropus boisei was first described in 1959 based on fossils from the Olduvai Gorge and now includes many fossils from Ethiopia to Malawi. Knowledge about its postcranial anatomy has remained elusive because, until recently, no postcranial remains could be reliably attributed to this taxon. Here, we report the first associated hand and upper limb skeleton (KNM-ER 47000) of P. boisei from 1.51 to 1.53 Ma sediments at Ileret, Kenya. While the fossils show a combination of primitive and derived traits, the overall anatomy is characterized by primitive traits that resemble those found in Australopithecus, including an oblique scapular spine, relatively long and curved ulna, lack of third metacarpal styloid process, gracile thumb metacarpal, and curved manual phalanges. Very thick cortical bone throughout the upper limb shows that P. boisei had great upper limb strength, supporting hypotheses that this species spent time climbing trees, although probably to a lesser extent than earlier australopiths. Hand anatomy shows that P. boisei, like earlier australopiths, was capable of the manual dexterity needed to create and use stone tools, but lacked the robust thumb of Homo erectus, which arguably reflects adaptations to the intensification of precision grips and tool use. KNM-ER 47000 provides conclusive evidence that early Pleistocene hominins diverged in postcranial and craniodental anatomy, supporting hypotheses of competitive displacement among these contemporaneous hominins.

Complete permanent mandibular dentition of early Homo from the upper Burgi Member of the Koobi Fora Formation, Ileret, Kenya.

The KNM-ER 64060 dentition derives from a horizon that most likely dates to between 2.02 and 2.03 Ma. A proximate series of postcranial bones (designated KNM-ER 64061) derives from the same siltstone unit and may be associated with the dentition, but their separation on the surface of the site leaves some room for doubt. KNM-ER 64060 is one of fewer than ten hominin specimens from the Early Pleistocene of East Africa that comprises a full or nearly complete mandibular dentition. Its taxonomic attribution is potentially significant, especially if the postcranial elements are related. At least three, and probably four hominin species, including Paranthropus boisei and Homo erectus (= H. ergaster), are known at about this time in East Africa. Other penecontemporaneous fossils have been referred to a single, highly variable species, H. habilis, or two taxa, namely H. habilis and H. rudolfensis. Although the weight of evidence supports the attribution of these specimens to two species, there is notable lack of agreement over the assignation of individual fossils. We take a conservative approach and group all such specimens under the designation "early Homo sp." for comparative purposes. KNM-ER 64060 is clearly attributable to Homo rather than Paranthropus. The preponderance of the evidence suggests that the affinities of KNM-ER 64060 are with fossils assigned to the early Homo sp. category rather than with H. erectus. This is indicated by the overall sizes of the KNM-ER 64060 canine, premolar and molar crowns, the size relationships of the P3 to P4, the relative narrowness of its premolar crowns, the cusp proportions of the M1 and especially those of the M2 and M3, and seemingly the possession of a two-rooted P4. Some of these comparisons suggest further that among the fossils comprising the early Homo sp. sample, the KNM-ER 64060 dentition exhibits greater overall similarity to specimens such as OH 7 and OH 16 that represent Homo habilis sensu stricto.

Humeral anatomy of the KNM-ER 47000 upper limb skeleton from Ileret, Kenya: Implications for taxonomic identification.

KNM-ER 47000 is a fossil hominin upper limb skeleton from the Koobi Fora Formation, Kenya (FwJj14E, Area 1A) that includes portions of the scapula, humerus, ulna, and hand. Dated to ∼1.52 Ma, the skeleton could potentially belong to one of multiple hominin species that have been documented in the Turkana Basin during this time, including Homo habilis, Homo erectus, and Paranthropus boisei. Although the skeleton lacks associated craniodental material, the partial humerus (described here) preserves anatomical regions (i.e., distal diaphysis, elbow joint) that are informative for taxonomic identification among early Pleistocene hominins. In this study, we analyze distal diaphyseal morphology and the shape of the elbow region to determine whether KNM-ER 47000 can be confidently attributed to a particular species. The morphology of the KNM-ER 47000 humerus (designated KNM-ER 47000B) is compared to that of other early Pleistocene hominin fossil humeri via the application of multivariate ordination techniques to both two-dimensional landmark data (diaphysis) and scale-free linear shape data (elbow). Distance metrics reflecting shape dissimilarity between KNM-ER 47000B and other fossils (and species average shapes) are assessed in the context of intraspecific variation within modern hominid species (Homo sapiens, Pan troglodytes, Gorilla gorilla, Pongo pygmaeus). Our comparative analyses strongly support attribution of KNM-ER 47000 to P. boisei. Compared to four other partial skeletons that have (justifiably or not) been attributed to P. boisei, KNM-ER 47000 provides the most complete picture of upper limb anatomy in a single individual. The taxonomic identification of KNM-ER 47000 makes the skeleton an important resource for testing future hypotheses related to P. boisei upper limb function and the taxonomy of isolated early Pleistocene hominin remains.

Inter-ray variation in metatarsal strength properties in humans and African apes: Implications for inferring bipedal biomechanics in the Olduvai Hominid 8 foot.

When measured as a ratio of mean midshaft diameter to bone length, the OH 8 fossil hominin foot exhibits a metatarsal (Mt) robusticity pattern of 1 > 5 > 3 > 4 > 2, which differs from the widely perceived "common" modern human pattern (1 > 5 > 4 > 3 > 2); African apes generally exhibit a third pattern (1 > 2 > 3 > 4 > 5). Largely because of the relative ranking of Mt2 and Mt5, OH 8 metatarsals structurally resemble the pattern exhibited by bipedal humans more than the pattern of quadrupedal and climbing African apes. Considering only these three phenotypes, however, discounts the potentially important functional implications of variation in modern human (and African ape) metatarsal robusticity patterns, suggesting that they are not useful for interpreting the specific biomechanics of a bipedal gait in fossils (i.e., whether it was modern human-like or not). Using computed tomography scans to quantify metatarsal midshaft cross-sectional geometry in a large sample of Homo (n=130), Gorilla (n=44) and Pan (n=80), we documented greater variation in metatarsal robusticity patterns than previously recognized in all three groups. While apes consistently show a 1 > 2 > 3 > 4 > 5 pattern in our larger sample, there does not appear to be a similarly precise single "common" human pattern. Rather, human metatarsals converge towards a 1 > 4/5 > 2/3 pattern, where metatarsals 4 and 5, and metatarsals 2 and 3, often "flip" positions relative to each other depending on the variable examined. After reassessing what a "common" human pattern could be based on a larger sample, the previously described OH 8 pattern of 1 > 5 > 3 > 4 > 2 is only observed in some humans (<6%) and almost never in apes (<0.5%). Although this suggests an overall greater similarity to (some) humans than to any ape in loading of the foot, the relatively rare frequency of these humans in our sample underscores potential differences in loading experienced by the medial and lateral columns of the OH 8 foot compared to modern humans.

Early Homo and the role of the genus in paleoanthropology.

The history of the discovery of early fossils attributed to the genus Homo has been contentious, with scholars disagreeing over the generic assignment of fossils proposed as members of our genus. In this manuscript I review the history of discovery and debate over early Homo and evaluate the various taxonomic hypotheses for the genus. To get a sense of how hominin taxonomy compares to taxonomic practice outside paleoanthropology, I compare the diversity of Homo to genera in other vertebrate clades. Finally, I propose a taxonomic model that hews closely to current models for hominin phylogeny and is consistent with taxonomic practice across evolutionary biology.

Pounding tools in HWK EE and EF-HR (Olduvai Gorge, Tanzania): Percussive activities in the Oldowan-Acheulean transition.

In this paper, we present pounded objects from excavations at HWK EE and EF-HR, which are studied from macro and microscopic perspectives. Analysis of HWK EE revealed one of the largest collections of percussive objects from Olduvai Gorge, while excavations at EF-HR have allowed us to recover a much wider collection of percussive tools than previously recorded. Differences are observed between the two localities. At the Acheulean site of EF-HR, percussive tools were predominantly used in the production of flakes and large cutting tools (LCTs). At the Oldowan site of HWK EE, the tool repertoire probably related to a wider range of activities, including bone breaking and bipolar knapping. Comparison of these two assemblages, potentially produced by different hominin species, helps provide a wider picture of pounding activities during the Oldowan-Acheulean transition at Olduvai Gorge.

Intra-individual metameric variation expressed at the enamel-dentine junction of lower post-canine dentition of South African fossil hominins and modern humans.

OBJECTIVES: The aim of this study is to compare the degree and patterning of inter- and intra-individual metameric variation in South African australopiths, early Homo and modern humans. Metameric variation likely reflects developmental and taxonomical issues, and could also be used to infer ecological and functional adaptations. However, its patterning along the early hominin postcanine dentition, particularly among South African fossil hominins, remains unexplored. MATERIALS AND METHODS: Using microfocus X-ray computed tomography (µXCT) and geometric morphometric tools, we studied the enamel-dentine junction (EDJ) morphology and we investigated the intra- and inter-individual EDJ metameric variation among eight australopiths and two early Homo specimens from South Africa, as well as 32 modern humans. RESULTS: Along post-canine dentition, shape changes between metameres represented by relative positions and height of dentine horns, outlines of the EDJ occlusal table are reported in modern and fossil taxa. Comparisons of EDJ mean shapes and multivariate analyses reveal substantial variation in the direction and magnitude of metameric shape changes among taxa, but some common trends can be found. In modern humans, both the direction and magnitude of metameric shape change show increased variability in M2 -M3 compared to M1 -M2 . Fossil specimens are clustered together showing similar magnitudes of shape change. Along M2 -M3 , the lengths of their metameric vectors are not as variable as those of modern humans, but they display considerable variability in the direction of shape change. CONCLUSION: The distalward increase of metameric variation along the modern human molar row is consistent with the odontogenetic models of molar row structure (inhibitory cascade model). Though much remains to be tested, the variable trends and magnitudes in metamerism in fossil hominins reported here, together with differences in the scale of shape change between modern humans and fossil hominins may provide valuable information regarding functional morphology and developmental processes in fossil species.

Australopithecus sediba and the emergence of Homo: Questionable evidence from the cranium of the juvenile holotype MH 1.

Malapa Hominin (MH) 1, an immature individual whose second permanent molars had recently reached occlusion at the time of death, is the holotype of Australopithecus sediba, a 2-myr-old South African taxon that has been hypothesized to link phylogenetically australopith-grade hominins to the Homo clade. Given the existence of 2.8 myr-old fossils of Homo in eastern Africa, this hypothesis implies a ghost lineage spanning at least 800 kyr. An alternative hypothesis posits a unique relationship between A. sediba and Australopithecus africanus, which predates the Malapa hominins in southern Africa and whose phylogenetic relationships remain ambiguous. The craniofacial morphology of MH 1 looms large in the framing of the two hypotheses. We evaluated these alternatives in two ways. First, we investigated whether the craniofacial morphology of MH 1 was ontogenetically stable at death. Based on data from a late-growth series of chimpanzee, gorilla, and modern human crania, we found that key aspects of MH 1's resemblance to Homo can be accounted for by its immaturity. Second, we studied MH 1 with an eye to identifying craniofacial synapomorphies shared with A. africanus. In this case, MH 1 shows unambiguous affinities in its zygomaticomaxillary and supraorbital morphology to crania from Sterkfontein Member 4, which we found to exhibit unusual derived morphology compared to Homo and other australopiths. We argue that MH 1 provides clear evidence that A. sediba was uniquely related to A. africanus and that the hypothesis of an extensive ghost lineage connecting A. sediba to the root of the Homo clade is unwarranted.

From Australopithecus to Homo: the transition that wasn't.

Although the transition from Australopithecus to Homo is usually thought of as a momentous transformation, the fossil record bearing on the origin and earliest evolution of Homo is virtually undocumented. As a result, the poles of the transition are frequently attached to taxa (e.g. A. afarensis, at ca 3.0 Ma versus H. habilis or H. erectus, at ca 2.0-1.7 Ma) in which substantial adaptive differences have accumulated over significant spans of independent evolution. Such comparisons, in which temporally remote and adaptively divergent species are used to identify a 'transition', lend credence to the idea that genera should be conceived at once as monophyletic clades and adaptively unified grades. However, when the problem is recast in terms of lineages, rather than taxa per se, the adaptive criterion becomes a problem of subjectively privileging 'key' characteristics from what is typically a stepwise pattern of acquisition of novel characters beginning in the basal representatives of a clade. This is the pattern inferred for species usually included in early Homo, including H. erectus, which has often been cast in the role as earliest humanlike hominin. A fresh look at brain size, hand morphology and earliest technology suggests that a number of key Homo attributes may already be present in generalized species of Australopithecus, and that adaptive distinctions in Homo are simply amplifications or extensions of ancient hominin trends.This article is part of the themed issue 'Major transitions in human evolution'.