Insights into the palaeobiology of an early Homo infant: multidisciplinary investigation of the GAR IVE hemi-mandible, Melka Kunture, Ethiopia.

Childhood is an ontogenetic stage unique to the modern human life history pattern. It enables the still dependent infants to achieve an extended rapid brain growth, slow somatic maturation, while benefitting from provisioning, transitional feeding, and protection from other group members. This tipping point in the evolution of human ontogeny likely emerged from early Homo. The GAR IVE hemi-mandible (1.8 Ma, Melka Kunture, Ethiopia) represents one of the rarely preserved early Homo infants (~ 3 years at death), recovered in a richly documented Oldowan archaeological context. Yet, based on the sole external inspection of its teeth, GAR IVE was diagnosed with a rare genetic disease-amelogenesis imperfecta (AI)-altering enamel. Since it may have impacted the child's survival, this diagnosis deserves deeper examination. Here, we reassess and refute this diagnosis and all associated interpretations, using an unprecedented multidisciplinary approach combining an in-depth analysis of GAR IVE (synchrotron imaging) and associated fauna. Some of the traits previously considered as diagnostic of AI can be better explained by normal growth or taphonomy, which calls for caution when diagnosing pathologies on fossils. We compare GAR IVE's dental development to other fossil hominins, and discuss the implications for the emergence of childhood in early Homo.

Assessing the status of the KNM-ER 42700 fossil using Homo erectus neurocranial development.

Based on ontogenetic data of endocranial shape, it has been proposed that a younger than previously assumed developmental status of the 1.5-Myr-old KNM-ER 42700 calvaria could explain why the calvaria of this fossil does not conform to the shape of other Homo erectus individuals. Here, we investigate (ecto)neurocranial ontogeny in H. erectus and assess the proposed juvenile status of this fossil using recent Homo sapiens, chimpanzees (Pan troglodytes), and Neanderthals (Homo neanderthalensis) to model and discuss changes in neurocranial shape from the juvenile to adult stages. We show that all four species share common patterns of developmental shape change resulting in a relatively lower cranial vault and expanded supraorbital torus at later developmental stages. This finding suggests that ectoneurocranial data from extant hominids can be used to model the ontogenetic trajectory for H. erectus, for which only one well-preserved very young individual is known. However, our study also reveals differences in the magnitudes and, to a lesser extent, directions of the species-specific trajectories that add to the overall shared pattern of neurocranial shape changes. We demonstrate that the very young H. erectus juvenile from Mojokerto together with subadult and adult H. erectus individuals cannot be accommodated within the pattern of the postnatal neurocranial trajectory for humans. Instead, the chimpanzee pattern might be a better 'fit' for H. erectus despite their more distant phylogenetic relatedness. The data are also compatible with an ontogenetic shape trajectory that is in some regards intermediate between that of recent H. sapiens and chimpanzees, implying a unique trajectory for H. erectus that combines elements of both extant species. Based on this new knowledge, neurocranial shape supports the assessment that KNM-ER 42700 is a young juvenile H. erectus if H. erectus followed an ontogenetic shape trajectory that was more similar to chimpanzees than humans.

Human uniqueness? Life history diversity among small-scale societies and chimpanzees.

BACKGROUND: Humans life histories have been described as "slow", patterned by slow growth, delayed maturity, and long life span. While it is known that human life history diverged from that of a recent common chimpanzee-human ancestor some ~4-8 mya, it is unclear how selection pressures led to these distinct traits. To provide insight, we compare wild chimpanzees and human subsistence societies in order to identify the age-specific vital rates that best explain fitness variation, selection pressures and species divergence. METHODS: We employ Life Table Response Experiments to quantify vital rate contributions to population growth rate differences. Although widespread in ecology, these methods have not been applied to human populations or to inform differences between humans and chimpanzees. We also estimate correlations between vital rate elasticities and life history traits to investigate differences in selection pressures and test several predictions based on life history theory. RESULTS: Chimpanzees' earlier maturity and higher adult mortality drive species differences in population growth, whereas infant mortality and fertility variation explain differences between human populations. Human fitness is decoupled from longevity by postreproductive survival, while chimpanzees forfeit higher potential lifetime fertility due to adult mortality attrition. Infant survival is often lower among humans, but lost fitness is recouped via short birth spacing and high peak fertility, thereby reducing selection on infant survival. Lastly, longevity and delayed maturity reduce selection on child survival, but among humans, recruitment selection is unexpectedly highest in longer-lived populations, which are also faster-growing due to high fertility. CONCLUSION: Humans differ from chimpanzees more because of delayed maturity and lower adult mortality than from differences in juvenile mortality or fertility. In both species, high child mortality reflects bet-hedging costs of quality/quantity tradeoffs borne by offspring, with high and variable child mortality likely regulating human population growth over evolutionary history. Positive correlations between survival and fertility among human subsistence populations leads to selection pressures in human subsistence societies that differ from those in modern populations undergoing demographic transition.

Molar crown formation times of fossil orangutan molars from Guangxi, China.

OBJECTIVES: We aimed to investigate molar enamel development in fossil orangutans from Guangxi and shed light on the evolution of Asian great apes. MATERIALS AND METHODS: We collected 32 fossil orangutan molars, most of which were from Guangxi apothecaries and the Guangxi Daxin Heidong cave, and prepared histological sections of each molar. We then characterized aspects of dental development, including long period line periodicity, number of Retzius lines and lateral enamel formation time, cuspal enamel thickness, and enamel formation time. RESULTS: The long period line periodicity in fossil orangutans ranged from 9 to 10 days (mean, 9.09 days). The molar lateral enamel formation time ranged from 1.48 to 3.17 years (540-1,152 days). Cuspal enamel thickness in fossil orangutan molars ranged from 949 to 2,535 µm, and cuspal enamel formation time ranged from 0.64 to 1.87 years. Molar enamel formation time of fossil orangutans ranged from 2.47 to 4.67 years. DISCUSSION: Long-period line periodicity of fossil orangutans from Guangxi was within the variation range of extant orangutans, and the average long period line periodicity (9.09 days) of fossil orangutans from Guangxi in this study was lower than the values for extant orangutans (9.5 days) and fossil orangutans (10.9 days) from Sumatra and Vietnam. Orangutan enamel thickness may have gradually decreased from the Middle Pleistocene to Holocene. Crown formation time of fossil orangutans was slightly longer than that of extant orangutans, and the M1 emergence age of fossil orangutans from Guangxi was about 4-6 years. These findings might indicate the regional difference or evolutionary changes in orangutans since Pleistocene. Dental development of the Guangxi fossil orangutans were more similar to that of Asian Miocene apes, suggesting the closer evolutionary relationship of orangutans to Miocene Asian fossil apes.

The character of parietal and orbital alterations in the superfamily Hominoidea (families Hominidae [exclusive of Homo] and Hylobotidae).

Parietal external surface disruption routinely referred to as porotic hyperostosis, and orbital alterations (cribra orbitalia), have been attributed to anemia-related bone marrow hyperplasia in humans. A recent study in humans identified that they were actually vascular in nature. Skeletons were examined and epi-illumination surface microscopy was performed on the parietal region and orbit of 156 Hominidae and 123 Hylobotidae to assess if these phenomena were trans-phylogenetic. Trans-cortical channels were recognized on the basis of visualized ectocranial surface defects penetrating the parietal; cribra orbitalia, by alteration of the normally smooth orbital roof appearance. Trans-cortical parietal channels, ranging in size from 20 to 100 µm, are rare in Gorilla and Pan troglodytes and absent in Pan paniscus. They are universally present in adult Pongo abeli and in Hylobatidae, independent of species. Cribra orbitalia was common in Hylobotidae, Pongo pygmaeus and P. abelii, less prevalent in adult P. troglodytes, and not recognized in any Gorilla gorilla or P. paniscus examined. The proliferative form predominated, with the exception of Hylobates concolor and muelleri, in which uncalcified vascular grooves predominated. No correlation was observed between the presence of either trans-cortical channels or cribra orbitalia and fractures, osteoarthritis, or inflammatory arthritis. Parietal alterations observed in apes are trans-cortical channels, analogous to those observed in humans, and do not represent porosity. Similarly, cribra orbitalia in apes is confirmed as vascular in nature. The proliferative form apparently represents calcification of blood vessel walls, indistinguishable from observations in humans. Predominant presence in adults rather than in juveniles suggests that both forms are acquired rather than developmental in derivation. Sex and bone alteration/disease-independence suggests that mechanical, endocrine, and inflammatory phenomena do not contribute to the development of either. Further, independent occurrence of trans-cortical channels and cribra orbitalia suggests that they do not have a shared etiology.

A comprehensive survey of Retzius periodicities in fossil hominins and great apes.

Recent studies have provided great insight into hominin life history evolution by utilizing incremental lines found in dental tissues to reconstruct and compare the growth records of extant and extinct humans versus other ape taxa. Among the hominins, studies that have examined Retzius periodicity (RP) variation have come to contradictory conclusions in some instances. To clarify RP variation among hominins and better place this variation in its broader evolutionary context, we conduct the most comprehensive analysis of published RP values for hominins and great apes to date. We gathered all available data from the literature on RP data from extant humans, great apes, and fossil hominins and assessed their variation using parametric and nonparametric analyses of variance. We also performed phylogenetic generalized least-squares regressions of RP data for these taxa as well as a larger set of hominoids for which RP data have been published against data for body mass, encephalization, and mean semicircular canal radius (a proxy for metabolic rate). Our results show that modern humans have a mean RP significantly differing from that of other hominins. Pongo also is significantly different from nearly all other taxa in all analyses. Our results also demonstrate that RP variation among hominins scales with respect to body mass, encephalization, and semicircular canal radius similarly to other hominids but that modern humans and Pongo stand out in this regard. Operating within the hypothesis that RP reflects autonomic biorhythms that regulate multiple life history variables, our results reinforce the idea that Homo sapiens has evolved a life history distinct from other hominins, even from other members of Homo, and suggest that many of these life history differences may be driven by hypothalamic output from the brain.

Utilizing auxology to understand ontogeny of extinct hominins: A case study on Homo naledi.

The methods used to study human growth and development (auxology) have not previously been applied within the setting of hominin maturation (ontogeny). Ontogeny is defined here as the pattern of biological change into an adult form, both at the individual and species level. The hominin fossil record has a lack of recovered immature materials, due to such factors as taphonomic processes that destroy pre-adults; the fragility of immature compared to adult bone; and the lower mortality rates of juveniles compared to adults. The recent discovery of pre-adult hominin skeletal material from a single, homogeneous Homo naledi species from the Rising Star cave system in South Africa provides the opportunity for a broader application of auxology methods and thus the need to understand their use in a modern context. Human auxology studies benefit from a robust database, across multiple populations, and with longitudinal studies in order to assess the patterns and variations in typical growth, development and life history stages. Here, we review the approach, vocabulary, and methods of these human studies, investigate commonalities in data with the fossil record, and then advance the reconstruction of ontogeny for the extinct hominin species H. naledi. To this end, we apply an auxology model into the paleontological context to broadly predict H. naledi birthweight of the offspring at 2.06 kg with a range (±1 SD) of 1.89 to 2.24 kg, with a length at birth 45.5 cm. We estimate a H. naledi juvenile partial skeleton DH7 to be a height of 111-125 cm at death.

Snapshots of human anatomy, locomotion, and behavior from Late Pleistocene footprints at Engare Sero, Tanzania.

Fossil hominin footprints preserve data on a remarkably short time scale compared to most other fossil evidence, offering snapshots of organisms in their immediate ecological and behavioral contexts. Here, we report on our excavations and analyses of more than 400 Late Pleistocene human footprints from Engare Sero, Tanzania. The site represents the largest assemblage of footprints currently known from the human fossil record in Africa. Speed estimates show that the trackways reflect both walking and running behaviors. Estimates of group composition suggest that these footprints were made by a mixed-sex and mixed-age group, but one that consisted of mostly adult females. One group of similarly-oriented trackways was attributed to 14 adult females who walked together at the same pace, with only two adult males and one juvenile accompanying them. In the context of modern ethnographic data, we suggest that these trackways may capture a unique snapshot of cooperative and sexually divided foraging behavior in Late Pleistocene humans.

Reassessment of the TM 1517 odonto-postcranial assemblage from Kromdraai B, South Africa, and the maturational pattern of Paranthropus robustus.

OBJECTIVES: The Pleistocene taxon Paranthropus robustus was established in 1938 following the discovery at Kromdraai B, South Africa, of the partial cranium TM 1517a and associated mandible TM 1517b. Shortly thereafter, a distal humerus (TM 1517g), a proximal ulna (TM 1517e), and a distal hallucial phalanx (TM 1517k) were collected nearby at the site, and were considered to be associated with the holotype. TM 1517a-b represents an immature individual; however, no analysis of the potentially associated postcranial elements has investigated the presence of any endostructural remnant of recent epiphyseal closure. This study aims at tentatively detecting such traces in the three postcranial specimens from Kromdraai B. MATERIALS AND METHODS: By using µXCT techniques, we assessed the developmental stage of the TM 1517b's C-M3 roots and investigated the inner structure of TM 1517g, TM 1517e, and TM 1517k. RESULTS: The M2 shows incompletely closed root apices and the M3 a half-completed root formation stage. The distal humerus was likely completely fused, while the proximal ulna and the distal hallucial phalanx preserve endosteal traces of the diaphyseo-epiphyseal fusion process. DISCUSSION: In the hominin fossil record, there are few unambiguously associated craniodental and postcranial remains sampling immature individuals, an essential condition for assessing the taxon-specific maturational patterns. Our findings corroborate the original association of the craniodental and postcranial remains representing the P. robustus type specimen. As with other Plio-Pleistocene hominins, the odonto-postcranial maturational pattern of TM 1517 more closely fits an African great ape rather than the extant human pattern.

Metameric variation of upper molars in hominoids and its implications for the diversification of molar morphogenesis.

Metameric variation of molar size is in part associated with the dietary adaptations of mammals and results from slight alterations of developmental processes. Humans and great apes exhibit conspicuous variation in tooth morphology both between taxa and across tooth types. However, the manner in which metameric variation in molars emerged among apes and humans via evolutionary alterations in developmental processes remains largely unknown. In this study, we compare the enamel-dentine junction of the upper molars of humans-which closely correlates with morphology of the outer enamel surface and is less affected by wear-with that of the other extant hominoids: chimpanzees, bonobos, gorillas, orangutans, and gibbons. We used the morphometric mapping method to quantify and visualize three-dimensional morphological variation, and applied multivariate statistical analyses. Results revealed the following: 1) extant hominoids other than humans share a common pattern of metameric variation characterized by a largely linear change in morphospace; this indicates a relatively simple graded change in metameric molar shape; 2) intertaxon morphological differences become less distinct from the mesial to distal molars; and 3) humans diverge from the extant ape pattern in exhibiting a distinct metameric shape change trajectory in the morphospace. The graded shape change and lower intertaxon resolution from the mesial to distal molars are consistent with the concept of a 'key' tooth. The common metameric pattern observed among the extant nonhuman hominoids indicates that developmental patterns underlying metameric variation were largely conserved during ape evolution. Furthermore, the human-specific metameric pattern suggests considerable developmental modifications in the human lineage.

Faster growth corresponds with shallower linear hypoplastic defects in great ape canines.

Deeper or more 'severe' linear enamel hypoplasia (LEH) defects are hypothesized to reflect more severe stress during development, but it is not yet clear how depth is influenced by intrinsic enamel growth patterns. Recent work documented inter- and intraspecific differences in LEH defect depth in extant great apes, with mountain gorillas having shallower defects than other taxa, and females having deeper defects than males. Here, we assess the correspondence of inter- and intraspecific defect depth and intrinsic aspects of enamel growth: enamel extension rates, outer enamel striae of Retzius angles, and linear enamel thickness. Thin sections of great ape canines (n = 40) from Gorilla beringei beringei, Gorilla gorilla gorilla, Pan troglodytes, and Pongo spp. were analyzed. Enamel extension rates were calculated within deciles of enamel-dentine junction length. Linear enamel thickness and the angle of intersection between striae of Retzius and the outer enamel surface were measured in the imbricational enamel. Mountain gorillas have faster enamel extension rates and shallower striae angles than the other taxa examined. Mountain gorillas have thinner imbricational enamel than western lowland gorillas and orangutans, but not chimpanzees. In the combined-taxon sample, females exhibit larger striae angles and thicker imbricational enamel than males. Enamel extension rates are highly negatively correlated with striae angles and LEH defect depth. Enamel growth variation corresponds with documented inter- and intraspecific differences in LEH defect depth in great ape canines. Mountain gorillas have shallower striae angles and faster extension rates than other taxa, which might explain their shallow LEH defect morphology and the underestimation of their LEH prevalence in previous studies. These results suggest that stressors of similar magnitude and timing might produce defects of different depths in one species or sex vs. another, which has implications for interpretations of stress histories in hominins with variable enamel growth patterns.

Endostructural morphology in hominoid mandibular third premolars: Discrete traits at the enamel-dentine junction.

The mandibular third premolar (P3) exhibits substantial differences in size and shape among hominoid taxa, and displays a number of discrete traits that have proven to be useful in studies of hominin taxonomy and phylogeny. Discrete traits at the enamel-dentine junction (EDJ) can be accurately assessed on moderately worn specimens, and often appear sharper than at the outer-enamel surface (OES). Here we use microtomography to image the P3 EDJ of a broad sample of extant apes, extinct hominins and modern humans (n = 100). We present typologies for three important premolar discrete traits at the EDJ (transverse crest, marginal ridge and buccal grooves), and score trait frequencies within our sample. We find that the transverse crest is variable in extant apes, while the majority of hominins display a transverse crest which runs directly between the two major premolar cusps. Some Neanderthals display a unique form in which the transverse crest fails to reach the protoconid. We find that mesial marginal ridge discontinuity is common in Australopithecus anamensis and Australopithecus afarensis while continuous marginal ridges largely characterize Australopithecus africanus and Paranthropus. Interrupted mesial and distal marginal ridges are again seen in Homo sapiens and Neanderthals. Premolar buccal grooves, previously identified at the OES as important for hominin systematics, are again found to show a number of taxon-specific patterns at the EDJ, including a clear difference between Australopithecus and Paranthropus specimens. However, their appearance may be dependent on the morphology of other parts of the crown such as the protoconid crest, and the presence of accessory dentine horns. Finally, we discuss rare variations in the form of dentine horns that underlie premolar cusps, and their potential homology to similar morphologies in other tooth positions.

Basicranial ontogeny comparison in Pan troglodytes and Homo sapiens and its use for developmental stage definition of KNM-ER 42700.

OBJECTIVE: This study aims to develop a comparative basis for assessing the developmental stage of KNM-ER 42700 based on the ontogenetic pattern of the ectocranial surface of the basicranium in modern humans and chimpanzees. MATERIALS AND METHODS: A total of 33 landmarks were collected from an ontogenetic sample of modern humans (80), chimpanzees (51), and 12 individuals classified as Homo erectus s.l. Ontogenetic trajectories were analyzed, and common aspects were extracted for the purpose of discriminating age groups. A regression of size on the extracted shape variables was used to investigate common ontogenetic allometry. RESULTS: The basicranial development of chimpanzees and humans follows different trajectories; however, similarities are also present. The common shape component of development extracted can be used to define age groups in both chimpanzees and modern humans. The extracted shape component presents a similar ontogenetic and static-allometric pattern in these two species. The developmental stages of H. erectus s.l. specimens were attributed following these common traits. Our analysis correctly assigned developmental stages to those specimens of H. erectus for which developmental ages are known. DISCUSSION: The component used for assessing the developmental stage has an ontogenetic allometric component. However, this shape component can discriminate age group irrespective of size and is no longer related to size when static allometry is considered. Adult H. erectus s.l. specimens were attributed to the adult category. KNM-WT 15000 fell with the late juvenile age group, whereas D2700 plotted in the region of overlap between the juvenile and adult age groups and Mojokerto with the younger age groups, as predicted by their known developmental ages. KNM-ER 42700 fell within the adult variability despite its incompletely fused spheno-occipital synchondrosis.

The deciduous dentition of Homo naledi: A comparative study.

In 2013, 2014 new hominin remains were uncovered in the Dinaledi chamber of the Rising Star cave system in South Africa. In 2015 Berger and colleagues identified these remains as belonging to a new species Homo naledi (Berger et al., 2015). Subsequent comparative studies of the skull, postcrania and permanent dentition have supported this taxonomic affiliation (Harcourt-Smith et al., 2015; Kivell et al., 2015; Irish et al., 2018). The deciduous teeth can offer unique insights into hominin evolution. Due to their early onset and rapid development their morphology is thought to be under stronger genetic control and less influenced by environment than are the permanent teeth. In this study we compared the H. naledi deciduous teeth from the 2013-2014 excavations to samples representing much of the hominin clade including Australopithecus afarensis, Australopithecus africanus, Paranthropus boisei, Paranthropus robustus, early Homo, Homo antecessor, Homo erectus s.l., Homo floresiensis, Middle Pleistocene Homo, Homo neanderthalensis, early Homo sapiens and recent H. sapiens from Sub-Saharan Africa. By making such a broad morphological comparison, we aimed to contextualize the Dinaledi hominins and to further assess the validity of their taxonomic assignment. Our analysis of the deciduous teeth revealed a unique combination of features that mirror (but also expand) that found in the permanent teeth. This mosaic includes an asymmetrical lower canine with a distal tubercle, an upper first molar with a large hypocone and epicrista associated with a mesial cuspule, a molarized lower first molar resembling Paranthropus, and upper and lower second molars that resemble later Homo in their lack of accessory cusps. The unique combination of deciduous dental characters supports previous studies assigning H. naledi to a new species, although its phylogenetic position vis-à-vis other Homo species remains ambiguous.

Enamel thickness and dental development in Rudapithecus hungaricus.

The fossil record of middle and late Miocene Eurasian hominoids has expanded considerably over the past few decades, particularly with the recovery of numerous isolated teeth and jaws. Scholars have turned to assessments of internal tooth structure and growth to make sense of the evolutionary radiations of these primates as well as their affinities to the living great apes (hominids). Here we characterize full-dentition enamel thickness and dental development in several juvenile Rudapithecus hungaricus individuals using multiple imaging modalities. Relative enamel thickness (RET) values for the anterior teeth and premolars of Rudapithecus are broadly akin to those of gorillas and chimpanzees and are thinner than those of orangutans. First molar RET values are most similar to chimpanzees, while posterior molar values are closer to thicker-enameled orangutans. When compared to Miocene hominoids, Rudapithecus shows an intermediate molar RET condition that is especially similar to other dryopithecines. Long-period line periodicity values are comparable to African apes and most Miocene hominoids, and lower than living and fossil orangutans. The mean cuspal daily secretion rate is similar to that of several other Miocene hominoids but is greater than extant great apes. Cusp-specific molar crown formation times generally exceed those of chimpanzees, are lower than those of orangutans, and are broadly like those of other Miocene apes. While Rudapithecus appears to have a somewhat unique pattern of enamel thickness and dental development relative to individual great ape genera, these structural and developmental features are consistent with its designation as a hominid.

Brain size growth in Australopithecus.

Postnatal growth is one of the proximate means by which humans attain massive adult brain size. Humans are characterized by the maintenance of prenatal brain growth rates into the first postnatal year, as well as an overall extended period of growth. The evolution of this pattern is difficult to assess due to its relatively brief duration and the underrepresentation of well-preserved fossil individuals who died during this short period. In this study, I use Monte Carlo methods to reconstruct postnatal brain growth rates in Australopithecus afarensis and Australopithecus africanus, based on estimates of neonatal brain size and of likely brain size and age at death of infant specimens (A.L. 333-105, DIK-1-1, and Taung). Neonatal brain size is reconstructed from the empirical scaling relationship among catarrhines which humans follow, and conservative estimates of fossils' chronological ages and brain sizes are drawn from the literature. Simulated distributions of these values are used to calculate average annual rates (ARs) of brain growth and proportional size change from birth (PSC), which are compared to resampled statistics from humans, chimpanzees and gorillas of known age and sex. Simulated ARs and PSCs for A. afarensis are significantly lower than those of chimpanzees and gorillas. Both ARs and PSCs for A. africanus are similar to chimpanzee and gorilla values. These results indicate that although these early hominins were derived in some aspects of brain anatomy, high rates of brain growth did not appear until later in human evolution. Moreover, findings also imply that brain growth rates are not a simple function of adult brain size. This study provides important new information about the evolution of brain growth, despite limitations inherent in fossil samples.

Safe Casting and Reliable Cusp Reconstruction Assisted by Micro-Computed Tomographic Scans of Fossil Teeth.

Dental replicas are frequently utilized in paleoanthropological studies of perikymata and enamel hypoplasia. However, fossil teeth are often fragile and worn, causing two problems: (1) the risk of damage by removing enamel fragments when impression-making material is separated from the fossil tooth surface, and (2) the need to reconstruct worn portions of the crown to assess perikymata number, distribution, and hypoplasia timing. This study presents the advantages of µCT data of canines and lateral incisors for (1) detecting cracks along the enamel-dentine junction (EDJ) which could cause damage when casting, and (2) reliably and non-destructively reconstructing worn or broken cusps. Fragile teeth of Homo naledi, Miocene, and Pleistocene specimens were µCT-scanned: 2D virtual sections and 3D models allowed for inspecting crack pattern beyond the external surface and 2D virtual sections were used to digitally reconstruct cusp tips (only Homo naledi). Micro-CT scans allowed cracks running along the EDJ and communicating with radial cracks in the enamel to be identified prior to casting. Cusp reconstructions using µCT data were conducted as precisely as when using thin-sections or photographs, and with high intra- and inter-observer agreement, while preserving the original specimen and affording numerous planes of virtual section. When available, µCT data should be inspected prior to tooth casting to exclude teeth that show a pattern of cracks that could lead to damage. Virtual sections allow for accessible, reliable, and non-destructive cusp reconstructions that may be used for developmental (e.g., perikymata and enamel hypoplasia) or enamel thickness studies. Anat Rec, 302:1516-1535, 2019. © 2018 American Association for Anatomy.

Developmental Loci Harbor Clusters of Accelerated Regions That Evolved Independently in Ape Lineages.

Some of the fastest evolving regions of the human genome are conserved noncoding elements with many human-specific DNA substitutions. These human accelerated regions (HARs) are enriched nearby regulatory genes, and several HARs function as developmental enhancers. To investigate if this evolutionary signature is unique to humans, we quantified evidence of accelerated substitutions in conserved genomic elements across multiple lineages and applied this approach simultaneously to the genomes of five apes: human, chimpanzee, gorilla, orangutan, and gibbon. We find roughly similar numbers and genomic distributions of lineage-specific accelerated regions (linARs) in all five apes. In particular, apes share an enrichment of linARs in regulatory DNA nearby genes involved in development, especially transcription factors and other regulators. Many developmental loci harbor clusters of nonoverlapping linARs from multiple apes, suggesting that accelerated evolution in each species affected distinct regulatory elements that control a shared set of developmental pathways. Our statistical tests distinguish between GC-biased and unbiased accelerated substitution rates, allowing us to quantify the roles of different evolutionary forces in creating linARs. We find evidence of GC-biased gene conversion in each ape, but unbiased acceleration consistent with positive selection or loss of constraint is more common in all five lineages. It therefore appears that similar evolutionary processes created independent accelerated regions in the genomes of different apes, and that these lineage-specific changes to conserved noncoding sequences may have differentially altered expression of a core set of developmental genes across ape evolution.

Patterns of lateral enamel growth in Homo naledi as assessed through perikymata distribution and number.

Perikymata, incremental growth lines visible on tooth enamel surfaces, differ in their distribution and number among hominin species, although with overlapping patterns. This study asks: (1) How does the distribution of perikymata along the lateral enamel surface of Homo naledi anterior teeth compare to that of other hominins? (2) When both perikymata distribution and number are analyzed together, how distinct is H. naledi from other hominins? A total of 19 permanent anterior teeth (incisors and canines) of H. naledi were compared, by tooth type, to permanent anterior teeth of other hominins: Australopithecus afarensis, Australopithecus africanus, Paranthropus robustus, Paranthropus boisei, Homo ergaster/Homo erectus, other early Homo, Neandertals, and modern humans, with varying sample sizes. Repeated measures analyses of the percentage of perikymata per decile of reconstructed crown height yielded several statistically significant differences between H. naledi and other hominins. Canonical variates analysis of percentage of perikymata in the cervical half of the crown together with perikymata number revealed that, in 8 of 19 cases, H. naledi teeth were significantly unlikely to be classified as other hominins, while exhibiting least difference from modern humans (especially southern Africans). In a cross-validated analysis, 68% of the H. naledi teeth were classified as such, while 32% were classified as modern human (most often southern African). Of 313 comparative teeth use for this analysis, only 1.9% were classified as H. naledi. What tends to differentiate H. naledi anterior tooth crowns from those of most other hominins, including some modern humans, is strongly skewed perikymata distributions combined with perikymata numbers that fall in the middle to lower ranges of hominin values. H. naledi therefore tends toward a particular combination of these features that is less often seen in other hominins. Implications of these data for the growth and development of H. naledi anterior teeth are considered.

The effect of ontogeny on estimates of KNM-WT 15000's adult body size.

The Homo erectus specimen KNM-WT 15000 has played a critical role in our understanding of body size evolution. New interpretations suggest that KNM-WT 15000 had a younger age-at-death and a more rapid ontogenetic trajectory than previously suggested. Recent fossil discoveries and new interpretations suggest a wide range of body size and shape variation in H. erectus. Based on these new insights, we argue that KNM-WT 15000's adult stature and body mass could have been much smaller than has been traditionally presented in the literature. Using chimpanzee and modern human growth trajectories, we bracketed the range of possibilities for KNM-WT 15000's adult body size between 160.0 and 177.7 cm (5'3″-5'10″) for stature and 60.0 and 82.7 kg (132-182 lbs.) for body mass. These estimates put KNM-WT 15000 near the mean rather than among the largest known H. erectus specimens.