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Background: Hybrids are expected to show greater phenotypic variation than their parental species, yet how hybrid phenotype expression varies with genetic distances in closely-related parental species remains surprisingly understudied. Here, we investigate pelage and morphometric trait variation in anthropogenic hybrids between four species of Brazilian Callithrix marmosets, a relatively recent primate radiation. Marmoset species are distinguishable by pelage phenotype and morphological specializations for eating tree exudates. In this work, we (1) describe qualitative phenotypic pelage differences between parental species and hybrids; (2) test whether significant quantitative differences exist between parental and hybrid morphometric phenotypes; and (3) determine which hybrid morphometic traits show heterosis, dysgenesis, trangression, or intermediacy relative to the parental trait. We investigated cranial and post-cranial morphometric traits, as most hybrid morphological studies focus on the former instead of the latter. Finally, we estimate mitogenomic distances between marmoset species from previously published data., Results: Marmoset hybrid facial and overall body pelage variation reflected novel combinations of coloration and patterns present in parental species. In morphometric traits, C. jacchus and C. penicillata were the most similar, while C. aurita was the most distinct, and C. geoffroyi trait measures fell between these species. Only three traits in C. jacchus x C. penicillata hybrids showed heterosis. We observed heterosis and dysgenesis in several traits of C. penicillata x C. geoffroyi hybrids. Transgressive segregation was observed in hybrids of C. aurita and the other species. These hybrids were also C. aurita-like for a number of traits, including body length. Genetic distance was closest between C. jacchus and C. penicillata and farthest between C. aurita and the other species., Conclusion: We attributed significant morphometric differences between marmoset species to variable levels of morphological specialization for exudivory in these species. Our results suggest that intermediate or parental species-like hybrid traits relative to the parental trait values are more likely in crosses between species with relatively lesser genetic distance. More extreme phenotypic variation is more likely in parental species with greater genetic distance, with transgressive traits appearing in hybrids of the most genetically distant parental species. We further suggest that fewer developmental disturbances can be expected in hybrids of more recently diverged parental species, and that future studies of hybrid phenotypic variation should investigate selective pressures on Callithrix cranial and post-cranial morphological traits., (© 2024. The Author(s).)

The Journal of Human Evolution (JHE) was founded 50 years ago when much of the foundation for how we think about human evolution was in place or being put in place, providing the main framework for how we consider our origins today. Here, we will explore historical developments, including early JHE outputs, as they relate to our understanding of the relationship between phenotypic variation and evolutionary process, and use that as a springboard for considering our current understanding of these links as applied to human evolution. We will focus specifically on how the study of variation itself has shifted us away from taxonomic and adaptationist perspectives toward a richer understanding of the processes shaping human evolutionary history, using literature searches and specific test cases to highlight this. We argue that natural selection, gene exchange, genetic drift, and mutation should not be considered individually when considering the production of hominin diversity. In this context, we offer suggestions for future research directions and reflect on this more complex understanding of human evolution and its broader relevance to society. Finally, we end by considering authorship demographics and practices in the last 50 years within JHE and how a shift in these demographics has the potential to reshape the science of human evolution going forward., Competing Interests: Declaration of competing interest The authors have no conflicts of interest to declare., (Copyright © 2022 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Previous scientific consensus saw human evolution as defined by adaptive differences (behavioural and/or biological) and the emergence of Homo sapiens as the ultimate replacement of non-modern groups by a modern, adaptively more competitive group. However, recent research has shown that the process underlying our origins was considerably more complex. While archaeological and fossil evidence suggests that behavioural complexity may not be confined to the modern human lineage, recent palaeogenomic work shows that gene flow between distinct lineages (for example, Neanderthals, Denisovans, early H. sapiens) occurred repeatedly in the late Pleistocene, probably contributing elements to our genetic make-up that might have been crucial to our success as a diverse, adaptable species. Following these advances, the prevailing human origins model has shifted from one of near-complete replacement to a more nuanced view of partial replacement with considerable reticulation. Here we provide a brief introduction to the current genetic evidence for hybridization among hominins, its prevalence in, and effects on, comparative mammal groups, and especially how it manifests in the skull. We then explore the degree to which cranial variation seen in the fossil record of late Pleistocene hominins from Western Eurasia corresponds with our current genetic and comparative data. We are especially interested in understanding the degree to which skeletal data can reflect admixture. Our findings indicate some correspondence between these different lines of evidence, flag individual fossils as possibly admixed, and suggest that different cranial regions may preserve hybridization signals differentially. We urge further studies of the phenotype to expand our ability to detect the ways in which migration, interaction and genetic exchange have shaped the human past, beyond what is currently visible with the lens of ancient DNA., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

Afro-Eurasian monkeys originated in the Miocene and are the most species-rich modern primate family. Molecular and fossil data have provided considerable insight into their evolutionary divergence, but we know considerably less about the evolutionary processes that underlie these differences. Here, we apply tests developed from quantitative genetics theory to a large (n > 3000) cranio-mandibular morphometric dataset, investigating the relative importance of adaptation (natural selection) and neutral processes (genetic drift) in shaping diversity at different taxonomic levels, an approach applied previously to monkeys of the Americas, apes, hominins, and other vertebrate taxa. Results indicate that natural selection, particularly for differences in size, plays a significant role in diversifying Afro-Eurasian monkeys as a whole. However, drift appears to better explain skull divergence within the subfamily Colobinae, and in particular the African colobine clade, likely due to habitat fragmentation. Small and declining population sizes make it likely that drift will continue in this taxon, with potentially dire implications for genetic diversity and future resilience in the face of environmental change. For the other taxa, many of whom also have decreasing populations and are threatened, understanding adaptive pressures similarly helps identify relative vulnerability and may assist with prioritising scarce conservation resources., (© 2022. The Author(s).)

Complex morphological traits are the product of many genes with transient or lasting developmental effects that interact in anatomical context. Mouse models are a key resource for disentangling such effects, because they offer myriad tools for manipulating the genome in a controlled environment. Unfortunately, phenotypic data are often obtained using laboratory-specific protocols, resulting in self-contained datasets that are difficult to relate to one another for larger scale analyses. To enable meta-analyses of morphological variation, particularly in the craniofacial complex and brain, we created MusMorph, a database of standardized mouse morphology data spanning numerous genotypes and developmental stages, including E10.5, E11.5, E14.5, E15.5, E18.5, and adulthood. To standardize data collection, we implemented an atlas-based phenotyping pipeline that combines techniques from image registration, deep learning, and morphometrics. Alongside stage-specific atlases, we provide aligned micro-computed tomography images, dense anatomical landmarks, and segmentations (if available) for each specimen (N = 10,056). Our workflow is open-source to encourage transparency and reproducible data collection. The MusMorph data and scripts are available on FaceBase ( www.facebase.org , https://doi.org/10.25550/3-HXMC ) and GitHub ( https://github.com/jaydevine/MusMorph )., (© 2022. The Author(s).)

The hypothesis that the evolution of humans involves hybridization between diverged species has been actively debated in recent years. We present the following novel evidence in support of this hypothesis: the analysis of nuclear pseudogenes of mtDNA ("NUMTs"). NUMTs are considered "mtDNA fossils" as they preserve sequences of ancient mtDNA and thus carry unique information about ancestral populations. Our comparison of a NUMT sequence shared by humans, chimpanzees, and gorillas with their mtDNAs implies that, around the time of divergence between humans and chimpanzees, our evolutionary history involved the interbreeding of individuals whose mtDNA had diverged as much as ~4.5 Myr prior. This large divergence suggests a distant interspecies hybridization. Additionally, analysis of two other NUMTs suggests that such events occur repeatedly. Our findings suggest a complex pattern of speciation in primate/human ancestors and provide one potential explanation for the mosaic nature of fossil morphology found at the emergence of the hominin lineage. A preliminary version of this manuscript was uploaded to the preprint server BioRxiv in 2017 (10.1101/134502).

The increasing awareness that hybridization, and resultant gene flow, plays a major role in animal diversification has led to a growing number of studies that have focused on assessing the morphological consequences of this process. Analyses of mammalian hybrids have identified skeletal effects of hybridization, including a suite of anomalous dental and sutural traits on the skull that are present at high frequencies in hybrid populations. These studies have also detected consistent patterns of morphological shape and size differences between hybrids and parental taxa across a wide variety of organisms. However, more research is required to understand the universality of these traits and shape/size differences. Building on these previous studies, a sample of genetically determined canid hybrids was examined, specifically the eastern coyote (Canis latrans var.), a hybrid between coyotes, wolves, and dogs, to test whether this group exhibits a comparable pattern of anomalous nonmetric characters, and to assess differences in craniomandibular shape and size. First, specimens of C. latrans var., C. latrans, and C. lupus were scored for anomalous traits, including supernumerary and rotated teeth, dental crowding, and sutural anomalies. Geometric morphometric analyses were then conducted on a subset of these individuals to explore craniomandibular size and shape variation, as well as allometry. The results are largely consistent with other studies, indicating that the incidence of dental anomalies, dental crowding, and sutural anomalies is significantly higher in hybrids. However, differences are not significant for supernumerary teeth. The exploration of morphometric variation identifies intermediate morphology in the hybrids, and some indication of greater morphological variability in the mandible. When these results are combined with previous studies, they suggest that skeletal signatures of hybridization are common to different mammalian taxa across multiple generations; however, some traits such as supernumerary teeth may be lost after a few generations., (© 2021 Wiley Periodicals LLC.)

Ancient DNA analyses have shown that interbreeding between hominin taxa occurred multiple times. Although admixture is often reflected in skeletal phenotype, the relationship between the two remains poorly understood, hampering interpretation of the hominin fossil record. Direct study of this relationship is often impossible due to the paucity of hominin fossils and difficulties retrieving ancient genetic material. Here, we use a sample of known ancestry hybrids between two closely related nonhuman primate taxa (Indian and Chinese Macaca mulatta) to investigate the effect of admixture on skeletal morphology. We focus on pelvic shape, which has potential fitness implications in hybrids, as mismatches between maternal pelvic and fetal cranial morphology are often fatal to mother and offspring. As the pelvis is also one of the skeletal regions that differs most between Homo sapiens and Neanderthals, investigating the pelvic consequences of interbreeding could be informative regarding the viability of their hybrids. We find that the effect of admixture in M. mulatta is small and proportional to the relatively small morphological difference between the parent taxa. Sexual dimorphism appears to be the main determinant of pelvic shape in M. mulatta. The lack of difference in pelvic shape between Chinese and Indian M. mulatta is in contrast to that between Neanderthals and H. sapiens, despite a similar split time (in generations) between the hybridizing pairs. Greater phenotypic divergence between hominins may relate to adaptations to disparate environments but may also highlight how the unique degree of cultural buffering in hominins allowed for greater neutral divergence. In contrast to some previous work identifying extreme morphologies in first- and second-generation hybrids, here the relationship between pelvic shape and admixture is linear. This linearity may be because most sampled animals have a multigenerational admixture history or because of relatively high constraints on the pelvis compared with other skeletal regions., Competing Interests: Conflict of interest statement The authors declare no conflict of interest., (Copyright © 2021 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

During the late Pleistocene, isolated lineages of hominins exchanged genes thus influencing genomic variation in humans in both the past and present. However, the dynamics of this genetic exchange and associated phenotypic consequences through time remain poorly understood. Gene exchange across divergent lineages can result in myriad outcomes arising from these dynamics and the environmental conditions under which it occurs. Here we draw from our collective research across various organisms, illustrating some of the ways in which gene exchange can structure genomic/phenotypic diversity within/among species. We present a range of examples relevant to questions about the evolution of hominins. These examples are not meant to be exhaustive, but rather illustrative of the diverse evolutionary causes/consequences of hybridization, highlighting potential drivers of human evolution in the context of hybridization including: influences on adaptive evolution, climate change, developmental systems, sex-differences in behavior, Haldane's rule and the large X-effect, and transgressive phenotypic variation., (© 2019 Wiley Periodicals, Inc.)

Hybridization occurs in a number of mammalian lineages, including among primate taxa. Analyses of ancient genomes have shown that hybridization between our lineage and other archaic hominins in Eurasia occurred numerous times in the past. However, we still have limited empirical data on what a hybrid skeleton looks like, or how to spot patterns of hybridization among fossils for which there are no genetic data. Here we use experimental mouse models to supplement previous studies of primates. We characterize size and shape variation in the cranium and mandible of three wild-derived inbred mouse strains and their first generation (F 1 ) hybrids. The three parent taxa in our analysis represent lineages that diverged over approximately the same period as the human/Neanderthal/Denisovan lineages and their hybrids are variably successful in the wild. Comparisons of body size, as quantified by long bone measurements, are also presented to determine whether the identified phenotypic effects of hybridization are localized to the cranium or represent overall body size changes. The results indicate that hybrid cranial and mandibular sizes, as well as limb length, exceed that of the parent taxa in all cases. All three F 1 hybrid crosses display similar patterns of size and form variation. These results are generally consistent with earlier studies on primates and other mammals, suggesting that the effects of hybridization may be similar across very different scenarios of hybridization, including different levels of hybrid fitness. This paper serves to supplement previous studies aimed at identifying F 1 hybrids in the fossil record and to introduce further research that will explore hybrid morphologies using mice as a proxy for better understanding hybridization in the hominin fossil record., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Recent fossil finds have highlighted extensive morphological diversity within our genus, Homo, and the co-existence of a number of species. However, little is known about the evolutionary processes responsible for producing this diversity. Understanding the action of these processes can provide insight into how and why our lineage evolved and diversified. Here, we examine cranial and mandibular variation and diversification from the earliest emergence of our genus at 2.8 Ma until the Late Pleistocene (0.126-0.0117 Ma), using statistical tests developed from quantitative genetics theory to evaluate whether stochastic (genetic drift) versus non-stochastic (selection) processes were responsible for the observed variation. Results show that random processes can account for species diversification for most traits, including neurocranial diversification, and across all time periods. Where selection was found to shape diversification, we show that: 1) adaptation was important in the earliest migration of Homo out of Africa; 2) selection played a role in shaping mandibular and maxillary diversity among Homo groups, possibly due to dietary differences; and 3) Homo rudolfensis is adaptively different from other early Homo taxa, including the earliest known Homo specimen. These results show that genetic drift, and, likely, small population sizes were important factors shaping the evolution of Homo and many of its novel traits, but that selection played an essential role in driving adaptation to new contexts., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

The data and methods presented in this article are supplementing the research article "Integration of mtDNA pseudogenes into the nuclear genome coincides with speciation of the human genus. A hypothesis", DOI: 10.1016/j.mito.2016.12.001 (Gunbin et al., 2017) [1]. Mitochondrial DNA is known to get inserted into nuclear DNA to form NUMTs, i.e. nuclear DNA pseudogenes of the mtDNA. We present here the sequences of selected NUMTs, in which time of integration can be determined with sufficient precision. We report their chromosomal positions , their position within the great ape mtDNA phylogeny, and their times of integration into the nuclear genome. The methods used to generate the data and to control their quality are also presented. The dataset is made publicly available to enable critical or extended analyzes.

Fragments of mitochondrial DNA are known to get inserted into nuclear DNA to form NUMTs, i.e. nuclear pseudogenes of the mtDNA. The insertion of a NUMT is a rare event. Hundreds of pseudogenes have been cataloged in the human genome. NUMTs are, in essence, a special type of mutation with their own internal timer, which is synchronized with an established molecular clock, the mtDNA. Thus insertion of NUMTs can be timed with respect to evolution milestones such as the emergence of new species. We asked whether NUMTs were inserted uniformly over time or preferentially during certain periods of evolution, as implied by the "punctuated evolution" model. To our surprise, the NUMT insertion times do appear nonrandom with at least one cluster positioned at around 2.8 million years ago (Ma). Interestingly, 2.8Ma closely corresponds to the time of emergence of the genus Homo, and to a well-documented period of major climate change ca. 2.9-2.5Ma. It is tempting to hypothesize that the insertion of NUMTs is related to the speciation process. NUMTs could be either "riders", i.e., their insertion could be facilitated by the overall higher genome rearrangement activity during speciation, or "drivers", i.e. they may more readily get fixed in the population due to positive selection associated with speciation. If correct, the hypothesis would support the idea that evolution of our genus may have happened in a rapid, punctuated manner., (Copyright © 2016 Elsevier B.V. and Mitochondria Research Society. All rights reserved.)

The species Homo naledi was recently named from specimens recovered from the Dinaledi Chamber of the Rising Star cave system in South Africa. This large skeletal sample lacks associated faunal material and currently does not have a known chronological context. In this paper, we present comprehensive descriptions and metric comparisons of the recovered cranial and mandibular material. We describe 41 elements attributed to Dinaledi Hominin (DH1-DH5) individuals and paratype U.W. 101-377, and 32 additional cranial fragments. The H. naledi material was compared to Plio-Pleistocene fossil hominins using qualitative and quantitative analyses including over 100 linear measurements and ratios. We find that the Dinaledi cranial sample represents an anatomically homogeneous population that expands the range of morphological variation attributable to the genus Homo. Despite a relatively small cranial capacity that is within the range of australopiths and a few specimens of early Homo, H. naledi shares cranial characters with species across the genus Homo, including Homo habilis, Homo rudolfensis, Homo erectus, and Middle Pleistocene Homo. These include aspects of cranial form, facial morphology, and mandibular anatomy. However, the skull of H. naledi is readily distinguishable from existing species of Homo in both qualitative and quantitative assessments. Since H. naledi is currently undated, we discuss the evolutionary implications of its cranial morphology in a range of chronological frameworks. Finally, we designate a sixth Dinaledi Hominin (DH6) individual based on a juvenile mandible., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

The discovery of Homo naledi has expanded the range of phenotypic variation in Homo, leading to new questions surrounding the mosaic nature of morphological evolution. Though currently undated, its unique morphological pattern and possible phylogenetic relationships to other hominin taxa suggest a complex evolutionary scenario. Here, we perform geometric morphometric analyses on H. naledi cranial and mandibular remains to investigate its morphological relationship with species of Homo and Australopithecus. We use Generalized Procrustes analysis to place H. naledi within the pattern of known hominin skull diversity, distributions of Procrustes distances among individuals to compare H. naledi and Homo erectus, and neighbor joining trees to investigate the potential phenetic relationships between groups. Our goal is to address a set of hypotheses relating to the uniqueness of H. naledi, its affinity with H. erectus, and the age of the fossils based on skull morphology. Our results indicate that, cranially, H. naledi aligns with members of the genus Homo, with closest affiliations to H. erectus. The mandibular results are less clear; H. naledi closely associates with a number of taxa, including some australopiths. However, results also show that although H. naledi shares similarities with H. erectus, some distances from this taxon - especially small-brained members of this taxon - are extreme. The neighbor joining trees place H. naledi firmly within Homo. The trees based on cranial morphology again indicate a close relationship between H. naledi and H. erectus, whereas the mandibular tree places H. naledi closer to basal Homo, suggesting a deeper antiquity. Altogether, these results emphasize the unique combination of features (H. erectus-like cranium, less derived mandible) defining H. naledi. Our results also highlight the variability within Homo, calling for a greater focus on the cause of this variability, and emphasizing the importance of using the total morphological package for species diagnoses., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Homo naledi is a recently discovered species of fossil hominin from South Africa. A considerable amount is already known about H. naledi but some important questions remain unanswered. Here we report a study that addressed two of them: "Where does H. naledi fit in the hominin evolutionary tree?" and "How old is it?" We used a large supermatrix of craniodental characters for both early and late hominin species and Bayesian phylogenetic techniques to carry out three analyses. First, we performed a dated Bayesian analysis to generate estimates of the evolutionary relationships of fossil hominins including H. naledi. Then we employed Bayes factor tests to compare the strength of support for hypotheses about the relationships of H. naledi suggested by the best-estimate trees. Lastly, we carried out a resampling analysis to assess the accuracy of the age estimate for H. naledi yielded by the dated Bayesian analysis. The analyses strongly supported the hypothesis that H. naledi forms a clade with the other Homo species and Australopithecus sediba. The analyses were more ambiguous regarding the position of H. naledi within the (Homo, Au. sediba) clade. A number of hypotheses were rejected, but several others were not. Based on the available craniodental data, Homo antecessor, Asian Homo erectus, Homo habilis, Homo floresiensis, Homo sapiens, and Au. sediba could all be the sister taxon of H. naledi. According to the dated Bayesian analysis, the most likely age for H. naledi is 912 ka. This age estimate was supported by the resampling analysis. Our findings have a number of implications. Most notably, they support the assignment of the new specimens to Homo, cast doubt on the claim that H. naledi is simply a variant of H. erectus, and suggest H. naledi is younger than has been previously proposed., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Hybridization is increasingly proving to be an important force shaping human evolution. Comparisons of both ancient and modern genomes have provided support for a complex evolutionary scenario over the past million years, with evidence for multiple incidents of gene exchange. However, to date, genetic evidence is still limited in its ability to pinpoint the precise time and place of ancient admixture. For that we must rely on evidence of admixture from the skeleton. The research presented here builds on previous work on the crania of baboon hybrids, focusing on the nasal cavity of olive baboons, yellow baboons, and first generation (F1) hybrids. The nasal cavity is a particularly important anatomical region for study, given the clear differentiation of this feature in Neanderthals relative to their contemporaries, and therefore it is a feature that will likely differ in a distinctive manner in hybrids of these taxa. Metric data consist of 45 linear, area, and volume measurements taken from CT scans of known-pedigree baboon crania. Results indicate that there is clear evidence for differences among the nasal cavities of the parental taxa and their F1 hybrids, including a greater degree of sexual dimorphism in the hybrids. There is also some evidence for transgressive phenotypes in individual F1 animals. The greatest amount of shape variation occurs in the anterior bony cavity, the choana, and the mid-nasopharynx. Extrapolating our results to the fossil record, we would expect F1 hybrid fossils to have larger nasal cavities, on average, than either parental taxon, with overall nasal cavity shape showing the most profound changes in regions that are distinct between the parental taxa (e.g., anterior nasal cavity). We also expect size and shape differences to be more pronounced in male F1 hybrids than in females. Because of pronounced anterior nasal cavity differences between Neanderthals and their contemporaries, we suggest that this model might be effective for examining the fossil record of late Pleistocene contact., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Homo naledi is a previously-unknown species of extinct hominin discovered within the Dinaledi Chamber of the Rising Star cave system, Cradle of Humankind, South Africa. This species is characterized by body mass and stature similar to small-bodied human populations but a small endocranial volume similar to australopiths. Cranial morphology of H. naledi is unique, but most similar to early Homo species including Homo erectus, Homo habilis or Homo rudolfensis. While primitive, the dentition is generally small and simple in occlusal morphology. H. naledi has humanlike manipulatory adaptations of the hand and wrist. It also exhibits a humanlike foot and lower limb. These humanlike aspects are contrasted in the postcrania with a more primitive or australopith-like trunk, shoulder, pelvis and proximal femur. Representing at least 15 individuals with most skeletal elements repeated multiple times, this is the largest assemblage of a single species of hominins yet discovered in Africa.

Plio-Pleistocene environmental change influenced the evolutionary history of many animal lineages in Africa, highlighting key roles for both climate and tectonics in the evolution of Africa's faunal diversity. Here, we explore diversification in the southern African chacma baboon Papio ursinus sensu lato and reveal a dominant role for increasingly arid landscapes during past glacial cycles in shaping contemporary genetic structure. Recent work on baboons (Papio spp.) supports complex lineage structuring with a dominant pulse of diversification occurring 1-2Ma, and yet the link to palaeoenvironmental change remains largely untested. Phylogeographic reconstruction based on mitochondrial DNA sequence data supports a scenario where chacma baboon populations were likely restricted to refugia during periods of regional cooling and drying through the Late Pleistocene. The two lineages of chacma baboon, ursinus and griseipes, are strongly geographically structured, and demographic reconstruction together with spatial analysis of genetic variation point to possible climate-driven isolating events where baboons may have retreated to more optimum conditions during cooler, drier periods. Our analysis highlights a period of continuous population growth beginning in the Middle to Late Pleistocene in both the ursinus and the PG2 griseipes lineages. All three clades identified in the study then enter a state of declining population size (Nef) through to the Holocene; this is particularly marked in the last 20,000 years, most likely coincident with the Last Glacial Maximum. The pattern recovered here conforms to expectations based on the dynamic regional climate trends in southern Africa through the Pleistocene and provides further support for complex patterns of diversification in the region's biodiversity.

Numerous studies suggest that the transition from Australopithecus to Homo was characterized by evolutionary innovation, resulting in the emergence and coexistence of a diversity of forms. However, the evolutionary processes necessary to drive such a transition have not been examined. Here, we apply statistical tests developed from quantitative evolutionary theory to assess whether morphological differences among late australopith and early Homo species in Africa have been shaped by natural selection. Where selection is demonstrated, we identify aspects of morphology that were most likely under selective pressure, and determine the nature (type, rate) of that selection. Results demonstrate that selection must be invoked to explain an Au. africanus-Au. sediba-Homo transition, while transitions from late australopiths to various early Homo species that exclude Au. sediba can be achieved through drift alone. Rate tests indicate that selection is largely directional, acting to rapidly differentiate these taxa. Reconstructions of patterns of directional selection needed to drive the Au. africanus-Au. sediba-Homo transition suggest that selection would have affected all regions of the skull. These results may indicate that an evolutionary path to Homo without Au. sediba is the simpler path and/or provide evidence that this pathway involved more reliance on cultural adaptations to cope with environmental change.

Hybridization may have played a substantial role in shaping the diversity of our evolving lineage. Although recent genomic evidence has shown that hybridization occurred between anatomically modern humans (AMHS) and Neanderthals, it remains difficult to pin down precisely where and when this gene flow took place. Investigations of the hybrid phenotype in primates and other mammals are providing models for identifying signatures of hybridization in the fossil record. However, our understanding of intra- and inter-taxon variation in hybrids is still limited. Moreover, there is little evidence from these studies that is pertinent to the question of how long hybrid skeletal traits persist in descendants, and therefore it is not clear whether observed hybrid phenotypes are evidence of recent (e.g., F1) or much earlier hybridization events. Here, we present an analysis updating a previous study of cranial variation in pedigreed olive and yellow baboons and their hybrids. Results suggest that traits previously associated with hybrids in baboons and other mammalian species are also present in this expanded data set; many of these traits are highly heritable, confirming a genetic basis for their variation in this mixed population. While F1 animals - and especially F1 males - still have the highest number of dental anomalies, these and other atypical traits persist into later hybrid generations (such as F2 and B1). Moreover, non-F1 recombinants also show extremely rare trait variations, including reduced canines and rotated teeth. However, these results must be considered in light of the possibility that some founding individuals may have themselves been unrecognized hybrids. Despite this, the data are compelling, and indicate once again that further controlled research remains to be done on primates and other mammals in order to better understand variation in the hybrid phenotype., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

The present report follows up on the findings of previous research, including recent bioarchaeological study of well-dated Khoesan skeletal remains, that posits long term biological continuity among the indigenous peoples of South Africa after the Pleistocene. The Arizona State University Dental Anthropology System was used to record key crown, root, and intraoral osseous nonmetric traits in six early-through-late Holocene samples from the Cape coasts. Based on these data, phenetic affinities and an identification of traits most important in driving intersample variation were determined using principal components analysis and the mean measure of divergence distance statistic. To expand biological affinity comparisons into more recent times, and thus preliminarily assess the dental impact of disproportionate non-Khoesan gene flow into local peoples, dental data from historic Khoekhoe and San were also included. Results from the prehistoric comparisons are supportive of population continuity, though a sample from Matjes River Rockshelter exhibits slight phenetic distance from other early samples. This and some insignificant regional divergence among these coastal samples may be related to environmental and cultural factors that drove low-level reproductive isolation. Finally, a close affinity of historic San to all samples, and a significant difference of Khoekhoe from most early samples is reflective of documented population history following immigration of Bantu-speakers and, later, Europeans into South Africa., (Copyright © 2014 Wiley Periodicals, Inc.)

Since the announcement of the species Australopithecus sediba, questions have been raised over whether the Malapa fossils represent a valid taxon or whether inadequate allowance was made for intraspecific variation, in particular with reference to the temporally and geographically proximate species Au. africanus. The morphology of mandibular remains of Au. sediba, including newly recovered material discussed here, shows that it is not merely a late-surviving morph of Au. africanus. Rather-as is seen elsewhere in the cranium, dentition, and postcranial skeleton-these mandibular remains share similarities with other australopiths but can be differentiated from the hypodigm of Au. africanus in both size and shape as well as in their ontogenetic growth trajectory.

SK 847 and StW 53 have often been cited as evidence for early Homo in South Africa. To examine whether midfacial morphology is in agreement with these attributions, we analyze Euclidean distances calculated from 3-D coordinates on the maxillae of SK 847 and StW 53, as well as Australopithecus africanus (Sts 5, Sts 71), Paranthropus robustus (SK 46, SK 48, SK 52, SK 83), early Homo (KNM-ER 1813, KNM-ER 1805, KNM-ER 3733, KNM-WT 15000), P. boisei (KNM-ER 406, KNM-WT 17000, KNM-WT 17400), Gorilla gorilla (n=116), Homo sapiens (n=342), Pan paniscus (n=21) and P. troglodytes (n=65). Multivariate analyses separate extant hominoids suggesting we have captured taxonomic affinity. With the exception of SK 847 and SK 52, South African fossils tend to cluster together. P. robustus differs substantially from East African robust megadonts. SK 847 and StW 53 resemble the East African Homo specimens that are the most australopith-like, such as KNM-WT 15000 and KNM-ER 1813. The resemblance between StW 53 and Homo is driven partly by similarities in maxillary size. When distances are scaled, StW 53 aligns with A. africanus, while SK 847 clusters primarily with early Homo., (Copyright © 2012 Elsevier GmbH. All rights reserved.)

Molecular studies have demonstrated a deep lineage split between the two gorilla species, as well as divisions within these taxa; estimates place this divergence in the mid-Pleistocene, with gene flow continuing until approximately 80,000 years ago. Here, we present analyses of skeletal data indicating the presence of substantial recent gene flow among gorillas at all taxonomic levels: between populations, subspecies, and species. Complementary analyses of DNA sequence variation suggest that low-level migration occurred primarily in a westerly-to-easterly direction. In western gorillas, the locations of hybrid phenotypes map closely to expectations based on population refugia and riverine barrier hypotheses, supporting the presence of significant vicariance-driven structuring and occasional admixture within this taxon. In eastern lowland gorillas, the high frequency of hybrid phenotypes is surprising, suggesting that this region represents a zone of introgression between eastern gorillas and migrants from the west, and underscoring the conservation priority of this critically endangered group. These results highlight the complex nature of evolutionary divergence in this genus, indicate that historical gene flow has played a major role in structuring gorilla diversity, and demonstrate that our understanding of the evolutionary processes responsible for shaping biodiversity can benefit immensely from consideration of morphological and molecular data in conjunction.

Modern baboons evolved as a distinct lineage prior to 2.5 Mya. Previous scenarios of diversification within this lineage have assessed the phylogenetic position of the chacma baboon of southern Africa relative to other baboons, but have not examined variation within this taxon. Here we provide a phylogenetic analysis of lineage diversity across the range of the chacma baboon, and show that: (1) chacma baboons diverged as a separate lineage at approximately 1.84 Mya; (2) the chacma lineage is characterised by a deep lineage split dividing chacmas into northeastern (1.52 Mya) and southwestern (1.22 Mya) clades; (3) ruacana baboons of Namibia form their own distinct monophyletic group within the southwestern clade, emerging approximately 0.68 Mya. These patterns likely result from a complex interplay of genetic drift and gene flow as the chacma lineage diversified across a broad geographic landscape during the climatically variable Plio-Pleistocene.

Various methodological approaches have been used for reconstructing fossil hominin remains in order to increase sample sizes and to better understand morphological variation. Among these, morphometric quantitative techniques for reconstruction are increasingly common. Here we compare the accuracy of three approaches--mean substitution, thin plate splines, and multiple linear regression--for estimating missing landmarks of damaged fossil specimens. Comparisons are made varying the number of missing landmarks, sample sizes, and the reference species of the population used to perform the estimation. The testing is performed on landmark data from individuals of Homo sapiens, Pan troglodytes and Gorilla gorilla, and nine hominin fossil specimens. Results suggest that when a small, same-species fossil reference sample is available to guide reconstructions, thin plate spline approaches perform best. However, if no such sample is available (or if the species of the damaged individual is uncertain), estimates of missing morphology based on a single individual (or even a small sample) of close taxonomic affinity are less accurate than those based on a large sample of individuals drawn from more distantly related extant populations using a technique (such as a regression method) able to leverage the information (e.g., variation/covariation patterning) contained in this large sample. Thin plate splines also show an unexpectedly large amount of error in estimating landmarks, especially over large areas. Recommendations are made for estimating missing landmarks under various scenarios., (Copyright 2009 Wiley-Liss, Inc.)

Three sympatric fossil cercopithecoid genera (Cercopithecoides, Parapapio, and Theropithecus) occur in Members 3 and 4 at the Makapansgat Limeworks hominin locality, South Africa, and their presence in a single ecosystem suggest a certain degree of ecological and/or dietary differentiation between taxa. Here, we explore the extent of dietary niche separation amongst these taxa using stable isotope (13C/12C, 18O/16O) and trace-element (Sr, Ba, Ca) analyses of fossil tooth enamel. In particular we searched for evidence of subtle niche separation between the more closely related, morphologically similar taxa of the genus Parapapio, as uncertainties exist around their taxonomy and taxonomic identification. Given these uncertainties, craniometric analyses were also performed to ground the dietary interpretations in a morphological context. The results found no clear taxonomic signal in the craniometric data for the Parapapio sample, and further indicate that this sample was no more variable morphologically than a single, geographically circumscribed, extant chacma baboon sample. In contrast, two overlapping dietary ecologies were found within this same Makapansgat Parapapio sample. Additionally, two widely differing dietary ecologies were found within the Cercopithecoides williamsi sample, while results for Theropithecus darti indicate a predominantly C4 diet. Hence, although biogeochemical dietary indicators point towards distinct dietary ecologies within and between fossil genera at Makapansgat, within the genus Parapapio disjunctions exist between the dietary categories and the taxonomic assignment of specimens., (Copyright 2007 Wiley-Liss, Inc.)

We assess craniometric variation in 153 individually dated human crania from South Africa with the aim of investigating genetic continuity/discontinuity during the Holocene. Evidence from the archaeological record is used to pinpoint likely episodes of genetic discontinuity. Craniometric data are then used to assess the likelihood of genetic change having occurred. Two periods of possible genetic discontinuity are identified: i) c. 4,000 BP, when an increase in overall population size, shifts in site organization and diet, and reduced mobility, were accompanied by reductions in stature; ii) c. 2,000 BP, when the herding of domesticates and the use of pottery vessels were introduced into the region. Results indicate that there was a decrease in cranial size and concomitant size-related changes in craniofacial shape between c.4,000 BP and 3,000 BP. This was followed almost immediately by a recovery in craniofacial size and a return to pre-4,000 BP craniofacial shape at c. 3,000 BP. This recovery continued gradually, extending into the herder period without any major shifts in morphology at 2,000 BP. It is suggested that the fluctuations in craniofacial size/shape were related to changes in environmental factors. Results obtained are consistent with long term continuity in South African Later Stone Age populations during the Holocene., ((c) 2007 Wiley-Liss, Inc.)

Buried in the same South African cave deposits as Australopithecus, fossil papionins have been referred to Parapapio (Pp. whitei, Pp. broomi, Pp. jonesi, Pp. antiquus), Papio (P. izodi, P. angusticeps, P. h. robinsoni), Theropithecus (e.g., T. darti), Gorgopithecus, or Dinopithecus on the basis of postcanine tooth size and descriptive morphology of the muzzle. The morphological patterns of variation that these papionins demonstrate can help to place the Australopithecus fossils into a biochronological context and provide valuable information for reconstructing regional Plio-Pleistocene turnover. To document these patterns of variation across fossil-bearing sites, we explore morphometric affinities within Parapapio, and between Parapapio and other Plio-Pleistocene taxa (Dinopithecus ingens, Papio angusticeps, Papio izodi, and Theropithecus darti) by analyzing a sample of interlandmark distances derived from 3-D coordinate data of the most complete fossil papionin specimens available. Bivariate and multivariate analyses show that Pp. whitei exhibits as much variation between sites and between individuals as Pp. broomi and Pp. whitei combined. Diversity in Parapapio at Makapansgat and Sterkfontein may suggest substantial time depth to the caves. Theropithecus darti, Dinopithecus ingens, Papio angusticeps, Pp. whitei from Bolt's Farm (BF 43), and Pp. jonesi from Sterkfontein (STS 565) differ considerably from one another. Other Parapapio specimens across sites form a separate cluster with P. izodi from Taung, suggesting a Pliocene age for this site., (Copyright 2006 Wiley-Liss, Inc.)

Many questions about developmental divergence in human (and non-human primate) evolution can be fruitfully explored through investigation of the extant primate phenotype. Here I discuss two approaches that use patterns of variation in extant primates to consider hypotheses of 'tinkering' both in their own lineages, and also as applied to the fossil record of human evolution. In the first, I show how comparisons of ontogenetic morphological integration in extant humans and apes can be used to consider the developmental underpinnings of the morphological change seen in the transition from the prognathic australopith face to the relatively smaller, orthognathic Homo face. In the second approach, I demonstrate how studies of craniofacial variation in hybrid baboons can be used as models for considering developmental divergence in Plio-Pleistocene primates, including fossil hominins. Of particular interest is the fact that unusual non-metric dental and sutural variation in these hybrids appears to be a sensitive indicator of evolutionary developmental divergence. Future studies would profit from focusing on the breadth and especially the overlap of morphological variation among extant primate taxa in order to determine the degree to which underlying genetic similarity in functional regions, and difference in regulatory regions, explains the variable primate phenotype.

Recent studies point to contact and possible admixture among contemporaneous hominin species during the Plio-Pleistocene. However, detection of hybridization in fossils-and especially fossil hominins-is contentious, and it is hindered in large part by our lack of understanding about how morphological hybridity is manifested in the primate skeleton. Here, we report on a study of known-pedigree, purebred yellow and olive baboons (n = 112) and their hybrids (n = 57), derived from the baboon colony of the Southwest Foundation for Biomedical Research. The hybrids were analyzed in two different groups: (1) F1 = olive x yellow first-generation hybrids; (2) B1 = olive x F1 backcross hybrids. Thirty-nine metric variables were tested for heterosis and dysgenesis. Nonmetric data were also collected from the crania. Results show that these primate hybrids are somewhat heterotic relative to their parental populations, are highly variable, and display novel phenotypes. These effects are most evident in the dentition and probably indicate the mixing of two separately coadapted genomes and the breakdown in the coordination of early development, despite the fact that these populations diverged fairly recently. Similar variation is also observed in museum samples drawn from natural hybrid zones. The results offer a strategy for detecting hybrid zones in the fossil record; implications for interpreting the hominin fossil record are discussed.

By investigating similarity in cranial covariation patterns, it is possible to locate underlying functional and developmental causes for the patterning, and to make inferences about the evolutionary forces that have acted to produce the patterns. Furthermore, establishing where these covariation patterns may diverge in ontogeny can offer insight into when selection may have acted on development. Here, covariation patterns are compared among adult and non-adult members of the African ape/human clade, in order to address three questions. First, are integration patterns constant among adult African apes and humans? Second, are they are constant in non-adults--i.e. throughout ontogeny? Third, if they are not constant, when do they diverge? Measurements are obtained from 677 crania of adult and non-adult African apes and humans. In order to address the first two questions, correlation matrices and theoretical integration matrices are compared using matrix correlation methods. The third question is evaluated by comparing correlation and variance/covariance patterns, using matrix correlation and random skewers methods, respectively, between adjacent age categories within each species, and between equivalent age categories among the four species. Results show that the hominoids share a similar pattern of ontogenetic integration, suggesting that common developmental/functional integrative processes may play an important role in keeping covariance structure stable across this lineage. However, there are some important differences in the magnitude of integration and in phenotypic covariance structure among the species, which may provide some insight into how selection acted to differentiate humans from the great apes.

Recent paleoanthropological discoveries reveal a diverse, potentially speciose human fossil record. Such extensive morphological diversity results from the action of divergent evolutionary forces on an evolving lineage. Here, we apply quantitative evolutionary theory to test whether random evolutionary processes alone can explain the morphological diversity seen among fossil australopith and early Homo crania from the Plio-Pleistocene. We show that although selection may have played an important role in diversifying hominin facial morphology in the late Pliocene, this is not the case during the early evolution of the genus Homo, where genetic drift was probably the primary force responsible for facial diversification.

Living species are often used as analogues for fossil ones. When this is done, the implicit assumption is made that hominids and living hominoids vary in the same way. This paper addresses the validity of this assumption by comparing patterns of facial variation among humans and African apes. In particular, it addresses three major questions that underlie approaches to reconstructing hominid relationships. First, is phenotypic variation similar between closely related species? Second, if it is dissimilar, why? Third, is it feasible to use analogue species for modeling purposes? Measurements are obtained from 542 crania of adult apes and humans. Care is taken to choose homologous data, and account for differences in population size and structure. Variance/covariance and correlation matrices among the species are compared using common principal component (CPC) analysis, random skewers methods and matrix correlations. Morphological distances (D(2)) are calculated between population means, and between randomized pairs of individuals within each population, to evaluate intraspecific variation. Morphological distances are also calculated between randomized pairs of individuals using the variation patterns of analogue populations, in order to evaluate the efficacy of such substitutions. Results show that while the hominoids share a similar pattern of facial variation overall, the patterns do diverge. This difference generally corresponds to the phylogenetic relationships among these species, suggesting that patterns of variation may have diverged through time in the large bodied hominoids. Because interpretation of relationships in the fossil record is confounded by a lack of understanding of how variation changes through time, exploration of such patterns of divergence can provide important clues to understanding human evolution. Additionally, neglecting to account for this divergence when using living analogues as variation "yardsticks" can give rise to interpretations of the fossil record that are more speciose than is warranted.

Recent evaluation of Neanderthal and modern human ontogeny suggests that taxon-specific features arose very early in development in both lineages, with early, possibly prenatal, morphological divergence followed by parallel postnatal developmental patterns. Here we use morphometric techniques to compare hominoid facial growth patterns, and show that this developmental phenomenon is, in fact, not unique to comparisons between Neanderthals and modern humans but extends to Australopithecus africanus and to the hominoid lineage more broadly. This finding suggests that a common pattern of juvenile facial development may be more widespread and that the roots of ontogenetically early developmental differentiation are deep-perhaps predating the ape/human split of 6+ million years ago., (Copyright 2002 Wiley-Liss, Inc.)

Quantitative genetic theory specifies evolutionary expectations for morphological diversification by genetic drift in a monophyletic clade. If genetic drift is responsible for the evolutionary morphological diversification of a clade, patterns of within- and between-taxon morphological variance/covariance should be proportional. We tested for proportionality of within- and between-species craniofacial morphological variation in 12 species of tamarins (genus Saguinus). We found that within- and between-taxon morphological variations across the entire genus were not proportional, and hence not likely to be due to genetic drift alone. The primary deviation from proportionality is that size and size-related shape in the cranium is more variable relative to other aspects of cranial morphology than expected under genetic drift, suggesting differential size selection between the two major clades, the small-bodied and large-bodied tamarins. Within each of these major clades, most of the interspecific variation is consistent with the pattern expected under genetic drift, although specific contrasts may indicate the involvement of differential selection. Morphological distances among taxa do not correspond very closely to the phylogeny derived from mtDNA. In particular, S. oedipus and S. geoffroyi are very distinct morphologically from the rest of the tamarins, although they are phylogenetically the sister clade to a clade containing S. midas and S. bicolor. Morphological similarity is not a good guide to phylogenetic affinity in the tamarins, especially with regard to deeper nodes in the phylogenetic tree., (Copyright 2002 Wiley-Liss, Inc.)

Constancy of variation/covariation structure among populations is frequently assumed in order to measure the differential selective forces which have caused population differentiation through evolutionary time. Following Steppan ([1997] Evolution 51:571-594), this assumption is examined among closely related tamarin species (genus Saguinus), using two distinct approaches applied to the task of evaluating similarity in patterns of morphological variation: common principal component analysis and matrix correlations. While the results of these analyses may appear contradictory, closer examination reveals them as complementary, highlighting the wisdom of combined methodologies. Overall, the results reveal a close relationship among the morphologically based variance structures of the tamarin species a relationship whose pattern is consistent with the pattern of phylogenetic relatedness as found via a molecular genetic study. More specifically, both methodological approaches provide some support for divergence of S. geoffroyi and S. oedipus (with regards to their patterns of morphological variation) from other tamarin species. This suggests that variance/covariance structure may have diverged through evolutionary time in the tamarin lineage, placing assumptions of constancy in doubt., (Copyright 2000 Wiley-Liss, Inc.)