Extant Primate Species Research Articles

Technical note: Micro-computed tomography calibration using dental tissue for bone mineral research.

Computed tomography (CT) and microcomputed tomography (μCT) require calibration against density phantoms scanned with specimens or during routine internal calibration for assessment of mineral concentration (MC) and density. In clinical studies involving bone, alternative calibration methods using bodily tissues and fluids ("phantomless" calibration) have been suggested. However, such tissues are seldom available in archeological and osteological research. This study investigates the potential of dental tissue as internal reference for calibration of μCT scans, facilitating the analysis of bone MC. We analyzed 70 molars from 24 extant primate species, including eight human teeth, each scanned with density phantoms for calibration. Our findings indicate that sampling specific regions of molars (lateral aspects of the mesial cusps) yields low variation in enamel and dentine MC values, averaging 1.27 g/cm3 (±0.03) for dentine and 2.25 g/cm3 (±0.03) for enamel. No significant differences were observed across molar types or amongscanning procedures, including scanner model, resolution, and filters. An ad hoc test on 12 mandibles revealed low variance in MC between the conventional phantom and dental tissue calibration methods; all 36 measurements (low, medium, and high MC for each mandible) were within 0.05 g/cm3 of each other -81% were within 0.03 g/cm3 and 94% within 0.04 g/cm3. Based on these results, we propose a new "phantomless" calibration technique using these mean enamel and dentine MC values. The presented phantomless calibration method could aid in the assessment of bone pathology and enhance the scope of studies investigating bone structure and physical property variations in archeological, osteological, and laboratory-based research.

Evolutionary Patterns of Intersexual Power

Dominance and leverage are both possible causes of social inequality. If sexual dimorphism influences patterns of intersexual dominance, we predicted that highly dimorphic species are constrained to exhibit male-biased power (MP), but species with low sexual dimorphism are free to demonstrate a broader range of intersexual power relationships. If market effects influence intersexual leverage, we predicted that females have more power when group composition is more male-biased and estrus is asynchronous. We analyzed data on intersexual power, sexual dimorphism, expected estrous overlap, and sex ratio for 79 extant primate species using phylogenetic logistic regression and ancestral state reconstructions. Although MP is more common, every major primate clade includes non-MP species. MP was associated with greater body mass and canine length dimorphism and with female-biased sex-ratios. Low estrous overlap was associated with non-MP. Although MP was reconstructed as likely ancestral for anthropoids, the last common ancestor of this clade probably did not exhibit high sexual dimorphism. The last common ancestor of catarrhines was probably highly dimorphic, potentially constraining intersexual power relationships. Non-MP probably evolved multiple times in primates and may be less common because multiple traits are linked to MP while fewer traits are associated with female-biased power or equality.

Homo sapiens and Neanderthals share high cerebral cortex integration into adulthood.

There is controversy around the mechanisms that guided the change in brain shape during the evolution of modern humans. It has long been held that different cortical areas evolved independently from each other to develop their unique functional specializations. However, some recent studies suggest that high integration between different cortical areas could facilitate the emergence of equally extreme, highly specialized brain functions. Here, we analyse the evolution of brain shape in primates using three-dimensional geometric morphometrics of endocasts. We aim to determine, firstly, whether modern humans present unique developmental patterns of covariation between brain cortical areas; and secondly, whether hominins experienced unusually high rates of evolution in brain covariation as compared to other primates. On the basis of analyses including modern humans and other extant great apes at different developmental stages, we first demonstrate that, unlike our closest living relatives, Homo sapiens retain high levels of covariation between cortical areas into adulthood. Among the other great apes, high levels of covariation are only found in immature individuals. Secondly, at the macro-evolutionary level, our analysis of 400 endocasts, representing 148 extant primate species and 6 fossil hominins, shows that strong covariation between different areas of the brain in H. sapiens and Homo neanderthalensis evolved under distinctly higher evolutionary rates than in any other primate, suggesting that natural selection favoured a greatly integrated brain in both species. These results hold when extinct species are excluded and allometric effects are accounted for. Our findings demonstrate that high covariation in the brain may have played a critical role in the evolution of unique cognitive capacities and complex behaviours in both modern humans and Neanderthals.

Current advances in primate genomics: novel approaches for understanding evolution and disease.

Primate genomics holds the key to understanding fundamental aspects of human evolution and disease. However, genetic diversity and functional genomics data sets are currently available for only a few of the more than 500 extant primate species. Concerted efforts are under way to characterize primate genomes, genetic polymorphism and divergence, and functional landscapes across the primate phylogeny. The resulting data sets will enable the connection of genotypes to phenotypes and provide new insight into aspects of the genetics of primate traits, including human diseases. In this Review, we describe the existing genome assemblies as well as genetic variation and functional genomic data sets. We highlight some of the challenges with sample acquisition. Finally,we explore how technological advances in single-cell functional genomics and induced pluripotent stem cell-derived organoids will facilitate our understanding of the molecular foundations of primate biology.

Testing for changes in rate of evolution and position of the climatic niche of clades

ABSTRACT There is solid recognition that phylogenetic effects must be acknowledged to appreciate climatic niche variability among species clades properly. Yet, most currently available methods either work at the intra‐specific level (hence they ignore phylogeny) or rely on the Brownian motion model of evolution to estimate phylogenetic effects on climatic niche variation. The Brownian motion model may be inappropriate to describe niche evolution in several cases, and even a significant phylogenetic signal in climatic variables does not indicate that the effect of shared ancestry was relevant to niche evolution. We introduce a new phylogenetic comparative method which describes significant changes in the width and position of the climatic niche at the inter‐specific (clade) level, while making no a priori assumption about how niche evolution took place. We devised the R function phylo.niche.shift to estimate whether the climatic niches of individual clades in the tree are either wider or narrower than expected, and whether the niche occupies unexpected climates. We tested phylo.niche.shift on realistic virtual species' distribution patterns applied to a phylogeny of 365 extant primate species. We demonstrate via simulations that the new method is fast and accurate under widely different climatic niche evolution scenarios. phylo.niche.shift showed that the capuchin monkeys and langurs occupy much wider, and prosimian much narrower, climatic niche space than expected by their phylogenetic positions. phylo.niche.shift may help to improve research on niche evolution by allowing researchers to test specific hypotheses on the factors affecting clades' realised niche width and position, and the potential effects of climate change on species' distributions.

A morphometric analysis of early Eocene Euprimate tarsals from Gujarat, India

Early Eocene primate postcranial bones from the Vastan lignite mine of Gujarat, India, have proven useful for understanding the haplorhine and strepsirrhine divergence. Previous analyses of material assigned to Asiadapidae supported interpretations that these primates were generalized arboreal quadrupeds, while the omomyid Vastanomys was likely to have been more proficient leaper than asiadapids. More recent examinations of long bone cross-sectional properties and calcaneal elongation have complicated the behavioral interpretations of these fossils. This study examines whole talar and calcaneal morphology of the Vastan material to refine the locomotor reconstructions of these fossils. A comparative sample of extant primate species representing various locomotor behaviors was obtained by accessing surface models from MorphoSource.org. Surface models of fossil specimens attributed to Asiadapis cambayensis, Marcgodinotius indicus, and Vastanomys major were generated from micro-computed tomography scans. A morphological analysis was carried out using weighted spherical harmonics, a Fourier-based method that represents surfaces using coefficients associated with a common set of spherical harmonic functions. The coefficients describing each surface were then used as shape variables in a principal components analysis. Significant differences between locomotor groups were assessed using nonparametric tests. Results from extant comparative samples show that locomotor behavior can be predicted from both talar and calcaneal morphology when phylogenetic relationships are known. Consistent with previous analyses, our results indicate that Asiadapis cambayensis and Marcgodinotius indicus were likely arboreal quadrupeds with some leaping capabilities. Vastanomys major is reconstructed as an arboreal quadruped with greater leaping proficiency than its asiadapid counterparts based on its talar morphology.

Nuku, a family of primate retrocopies derived from KU70.

The gene encoding the ubiquitous DNA repair protein, Ku70p, has undergone extensive copy number expansion during primate evolution. Gene duplications of KU70 have the hallmark of long interspersed element-1 mediated retrotransposition with evidence of target-site duplications, the poly-A tails, and the absence of introns. Evolutionary analysis of this expanded family of KU70-derived “NUKU” retrocopies reveals that these genes are both ancient and also actively being created in extant primate species. NUKU retrocopies show evidence of functional divergence away from KU70, as evinced by their altered pattern of tissue expression and possible tissue-specific translation. Molecular modeling predicted that amino acid changes in Nuku2p at the interaction interface with Ku80p would prevent the assembly of the Ku heterodimer. The lack of Nuku2p-Ku80p interaction was confirmed by yeast two-hybrid assay, which contrasts the robust interaction of Ku70p-Ku80p. While several NUKU retrocopies appear to have been degraded by mutation, NUKU2 shows evidence of positive natural selection, suggesting that this retrocopy is undergoing neofunctionalization. Although Nuku proteins do not appear to antagonize retrovirus transduction in cell culture, the observed expansion and rapid evolution of NUKUs could be being driven by alternative selective pressures related to infectious disease or an undefined role in primate physiology.

Molecular evolutionary analysis of human primary microcephaly genes

BackgroundThere has been a rapid increase in the brain size relative to body size during mammalian evolutionary history. In particular, the enlarged and globular brain is the most distinctive anatomical feature of modern humans that set us apart from other extinct and extant primate species. Genetic basis of large brain size in modern humans has largely remained enigmatic. Genes associated with the pathological reduction of brain size (primary microcephaly-MCPH) have the characteristics and functions to be considered ideal candidates to unravel the genetic basis of evolutionary enlargement of human brain size. For instance, the brain size of microcephaly patients is similar to the brain size of Pan troglodyte and the very early hominids like the Sahelanthropus tchadensis and Australopithecus afarensis.ResultsThe present study investigates the molecular evolutionary history of subset of autosomal recessive primary microcephaly (MCPH) genes; CEP135, ZNF335, PHC1, SASS6, CDK6, MFSD2A, CIT, and KIF14 across 48 mammalian species. Codon based substitutions site analysis indicated that ZNF335, SASS6, CIT, and KIF14 have experienced positive selection in eutherian evolutionary history. Estimation of divergent selection pressure revealed that almost all of the MCPH genes analyzed in the present study have maintained their functions throughout the history of placental mammals. Contrary to our expectations, human-specific adoptive evolution was not detected for any of the MCPH genes analyzed in the present study.ConclusionBased on these data it can be inferred that protein-coding sequence of MCPH genes might not be the sole determinant of increase in relative brain size during primate evolutionary history.

Morphological integration of the canine region within the hominine alveolar arch

The early hominin record is characterized by numerous shifts in dental proportions (e.g., canine reduction and megadontia) linked to changes in diet and social behavior. Recent studies suggest that hominins exhibit a reduction in the magnitude of covariation between the anterior and posterior dental components compared with other extant great apes. They point toward, but do not directly test, the relative independence of canine morphology within the hominin alveolar arch. This study focuses specifically on the how the canine region covaries with other regions of the dental arch because the canine region has drastically reduced in size and changed in shape across human evolution. We examine extant primate species most commonly used as a comparative framework for fossil hominin morphology: Gorilla gorilla (n = 27), Pan troglodytes (n = 27), and Homo sapiens (n = 30). We used geometric morphometric methods to test for size and shape covariation between the canine region with other dental regions. We also examined the influence of sexual dimorphism and allometry on intraspecific and interspecific patterns of covariation. The analysis of size and shape covariation between the mandibular canine and other individual tooth regions elucidated complex, species-specific, and sex-specific morphological relationships in the mandibular alveolar arch. There was little evidence to support different patterns of morphological integration between humans on the one hand and nonhuman apes on the other. Canine region morphology was relatively independent from other dental regions across species based on shape and did not significantly covary more with either the incisor or postcanine region in any species. The size correlations between the canine and other dental regions were moderate to high. The species-specific results of this study question the ability to make a priori assumptions about morphological integration in the extant hominin mandibular alveolar arch and its application to the fossil record.

Daily Distance Traveled Is Associated with Greater Brain Size in Primates

Explanations for the brain size increments through primate and, particularly, human evolution are numerous. Commonly, these hypotheses rely on the influence that behavioral and ecological variables have on brain size in extant primates, such as diet quality, social group size, or home range (HR) area. However, HR area does not reflect the time spent moving. As such, it has not been properly addressed whether the effort involved in movement could have affected brain size evolution in primates. This study aimed to test the influence of daily movement on primates’ brain sizes, controlling for these other behavioral and ecological factors. We used a large comparative dataset of extant primate species and phylogenetic comparative methods. Our results show a significant correlation between daily movement and brain mass, which is not explained by the influence of diet, social group size, HR, or body mass. Hence, from an evolutionary timescale, a longer daily movement distance is not a constraining factor for the energetic investment in a larger brain. On the contrary, increased mobility could have contributed to brain mass incrementations through evolution.

Current and future trends in socio-economic, demographic and governance factors affecting global primate conservation.

Currently, ~65% of extant primate species (ca 512 species) distributed in 91 countries in the Neotropics, mainland Africa, Madagascar, South Asia and Southeast Asia are threatened with extinction and 75% have declining populations as a result of deforestation and habitat loss resulting from increasing global market demands, and land conversion for industrial agriculture, cattle production and natural resource extraction. Other pressures that negatively impact primates are unsustainable bushmeat hunting, the illegal trade of primates as pets and as body parts, expanding road networks in previously isolated areas, zoonotic disease transmission and climate change. Here we examine current and future trends in several socio-economic factors directly or indirectly affecting primates to further our understanding of the interdependent relationship between human well-being, sustainable development, and primate population persistence. We found that between 2001 and 2018 ca 191 Mha of tropical forest (30% canopy cover) were lost as a result of human activities in the five primate range regions. Forty-six percent of this loss was in the Neotropics (Mexico, Central and South America), 30% in Southeast Asia, 21% in mainland Africa, 2% in Madagascar and 1% in South Asia. Countries with the greatest losses (ca 57% of total tree cover loss) were Brazil, Indonesia, DRC, China, and Malaysia. Together these countries harbor almost 50% of all extant primate species. In 2018, the world human population was estimated at ca 8bn people, ca 60% of which were found in primate range countries. Projections to 2050 and to 2100 indicate continued rapid growth of the human populations in these five primate range regions, with Africa surpassing all the other regions and totaling ca 4bn people by the year 2100. Socioeconomic indicators show that, compared to developed nations, most primate range countries are characterized by high levels of poverty and income inequality, low human development, low food security, high levels of corruption and weak governance. Models of Shared Socioeconomic Pathway scenarios (SSPs) projected to 2050 and 2100 showed that whereas practices of increasing inequality (SSP4) or unconstrained growth in economic output and energy use (SSP5) are projected to have dire consequences for human well-being and primate survivorship, practices of sustainability-focused growth and equality (SSP1) are expected to have a positive effect on maintaining biodiversity, protecting environments, and improving the human condition. These results stress that improving the well-being, health, and security of the current and future human populations in primate range countries are of paramount importance if we are to move forward with effective policies to protect the world’s primate species and promote biodiversity conservation.

Broad-scale morpho-functional traits of the mandible suggest no hard food adaptation in the hominin lineage

An on-going debate concerning the dietary adaptations of archaic hominins and early Homo has been fuelled by contradictory inferences obtained using different methodologies. This work presents an extensive comparative sample of 30 extant primate species that was assembled to perform a morpho-functional comparison of these taxa with 12 models corresponding to eight fossil hominin species. Finite Element Analysis and Geometric Morphometrics were employed to analyse chewing biomechanics and mandible morphology to, firstly, establish the variation of this clade, secondly, relate stress and shape variables, and finally, to classify fossil individuals into broad ingesta related hardness categories using a support vector machine algorithm. Our results suggest that some hominins previously assigned as hard food consumers (e.g. the members of the Paranthropus clade) in fact seem to rely more strongly on soft foods, which is consistent with most recent studies using either microwear or stable isotope analyses. By analysing morphometric and stress results in the context of the comparative framework, we conclude that in the hominin clade there were probably no hard-food specialists. Nonetheless, the biomechanical ability to comminute harder items, if required as fallback option, adds to their strategy of increased flexibility.

Increased terrestriality in a Neotropical primate living on islands with reduced predation risk

An arboreal lifestyle is thought to be central to primate origins, and most extant primate species still live in the trees. Nonetheless, terrestrial locomotion is a widespread adaptation that has arisen repeatedly within the primate lineage. The absence of terrestriality among the New World monkeys (Platyrrhini) is thus notable and raises questions about the ecological pressures that constrain the expansion of platyrrhines into terrestrial niches. Here, we report the results of a natural experiment, comparing patterns of terrestrial behavior in white-faced capuchin monkeys (Cebus capucinus imitator) living on two islands off the Pacific coast of Panama that lack mammalian predators (island sites) with the behavior of capuchins at three sites in central Panama with more intact predator communities (mainland sites). Surveys with camera traps revealed increased terrestriality in island vs. mainland sites. Capuchin detection rates were higher, the range of party sizes observed was larger, and individuals engaged in a wider range of terrestrial behaviors on the islands lacking mammalian predators. Furthermore, females carrying infants were frequently photographed on the ground at the island sites, but never at the mainland sites. These findings support the long-standing hypothesis that predators constrain the exploitation of terrestrial niches by primates. These results are also consistent with the hypothesis that arboreal locomotion imposes costs that primates will avoid by walking on the ground when predation risk is low.

A Morphometric Analysis of Early Eocene Euprimate Calcanei from Gujarat, India

Postcrania dating back to the early Eocene from the Cambay Shale Formation of Gujarat, India, provide a unique window for understanding early primate behavior. Previous metric analyses of postcrania assigned to Asiadapinae indicate that these primates were generalized arboreal quadrupeds. Examination of long bone cross‐sectional properties, however, complicates the initial behavioral interpretations because it shows that these fossils overlap with primates of differing locomotor behaviors. The purpose of this study is to re‐examine calcaneal morphology of the Indian material in order to refine the locomotor reconstructions of these fossils.Surface models of fossil calcanei (n=6) were generated from micro‐computed tomography scans while those (n=73) representing 23 extant primate species were downloaded from MorphoSource. A weighted spherical harmonics analysis was carried out on the total sample. This morphological analysis represents genus‐zero surfaces using a common set of spherical harmonic functions. The harmonic coefficients associated with each calcaneus serve as shape descriptors that can be summarized via principal components (PC) analysis. Bivariate plots and models representing plus and minus two standard deviations of each PC were created to visualize morphological differences.The first three PCs explain 52.4%, 9.7%, and 9.3% of the morphological variation in the sample respectively (71.4% combined). The first PC differentiates primates with elongated calcanei that are more likely to engage in vertical clinging and leaping, from more quadrupedal primates. The second PC distinguishes calcanei of slow climbers from primates that engage in other forms of locomotion (e.g., leaping and quadrupedalism). Extant primate morphology seems to follow a continuum in which relative anteroposterior calcaneal elongation negatively correlates with slow climber calcaneal morphology. As expected, the calcaneal morphology of Marcgodinotius indicus aligns with that of extant lemuriforms, and most closely resembles that of a generalized arboreal quadruped with some leaping ability. There are two fossils tentatively attributed to Asiadapis cambayensis included in the analysis. One groups with other generalized arboreal quadrupeds, while the other falls outside the range of calcaneal variation observed in extant primates. The outlier does not follow the general pattern of calcaneal morphology observed in extant primates; instead it is characterized by some aspects of slow climber morphology, but with a relative anteroposterior calcaneal elongation. These results may reflect the generalized morphology of this fossil, and consequently the behavior of this primitive euprimate.Support or Funding InformationSupported by NSF DGE‐1746891, National Geographic Society, Leakey Foundation

Mandibular symphyseal fusion in fossil primates: Insights from correlated patterns of jaw shape and masticatory function in living primates

Variation in primate masticatory form and function has been extensively researched through both morphological and experimental studies. As a result, symphyseal fusion in different primate clades has been linked to either the recruitment of vertically directed balancing-side muscle force, the timing and recruitment of transversely directed forces, or both. This study investigates the relationship between jaw muscle activity patterns and morphology in extant primates to make inferences about masticatory function in extinct primates, with implications for understanding the evolution of symphyseal fusion. Three-dimensional mandibular landmark data were collected for 31 extant primates and nine fossil anthropoids and subfossil lemur species. Published electromyography (EMG) data were available for nine of the extant primate species. Partial least squares analysis and phylogenetic partial least squares analysis were used to identify relationships between EMG and jaw shape data and evaluate variation in jaw morphology. Primates with partial and complete symphyseal fusion exhibit shape-function patterns associated with the wishboning motor pattern and loading regime, in contrast to shape-function patterns of primates with unfused jaws. All fossil primates examined (except Apidium) exhibit jaw morphologies suggestive of the wishboning motor pattern demonstrated in living anthropoids and indriids. Partial fusion in Catopithecus, similar to indriids and some subfossil lemurs, may be sufficient to resist, or transfer, some amounts of transversely directed balancing-side muscle force at the symphysis, representing a transition to greater reliance on transverse jaw movement during mastication. Furthermore, possible functional convergences in physiological patterns during chewing (i.e., Archaeolemur) are identified.

Macroevolutionary effects on primate trophic evolution and their implications for reconstructing primate origins

The visual-predation hypothesis proposes that certain derived features shared by crown primates reflect an insectivorous ancestry. Critics of this idea have argued that because insectivory is uncommon among extant primates it is unlikely to have been a major influence on early primate evolution. According to this perspective, the low frequency of insectivory indicates that it is an apomorphic deviation from the mostly conserved primate ecological pattern of herbivory. The present study tests two alternative hypotheses that are compatible with an insectivorous ancestor: (1) that trophic evolution was biased, such that herbivory evolved repeatedly with few shifts back to insectivory, and (2) that insectivorous lineages have diversified at a lower rate than herbivorous lineages owing to differential trophic effects on speciation and extinction probabilities. Model-based analysis conducted using trait data for 307 extant primate species indicates that rates of transition into and out of insectivory are similar, rejecting the hypothesis of biased trophic evolution. On the other hand, the hypothesis of asymmetric diversification is supported, with insectivorous lineages having a lower rate of diversification than herbivorous lineages. This correlation is mediated by activity pattern: insectivory occurs mostly in nocturnal lineages, which have a lower diversification rate than diurnal lineages. The frequency of insectivory also appears to have been shaped by repeated transitions into ecological contexts in which insectivory is absent (large body size) or rare (diurnality). These findings suggest that the current distribution of trophic strategies among extant primates is the result of macroevolutionary processes that have favored the proliferation and persistence of herbivory relative to insectivory. This conclusion implies that the low frequency of insectivory is not necessarily evidence against the visual-predation hypothesis.

The primate extinction crisis in China: immediate challenges and a way forward

China is facing an unprecedented set of challenges in balancing the effects of economic development and global climate change with environmental protection and maintaining biodiversity. Although positive steps have been undertaken to remedy this situation, currently 80% of China’s 25 extant primate species are threatened, 15–18 species have population sizes of less than 3000 individuals, and two species of gibbons and one species of langur have been extirpated over the past few decades. Today, virtually all species of primates in China inhabit fragmented landscapes and are distributed in small isolated subpopulations with limited opportunities to exchange individuals or genetic information. Here we present a historical framework examining how human-induced environmental changes, particularly since the second half of the 20th century, accelerated primate population decline in China. In addition, we modeled the expected spatial conflict between agricultural expansion and primate distributions over the next 25–75 years and assessed the current overlap between protected areas and primate distributions. Depending on the assumptions of the spatial conflict model, primate distributions are expected to decline by an additional 51–87% by the year 2100. Thus, unless large-scale conservation policies are implemented immediately the current trend of primate population decline, local extirpation, and species extinctions will accelerate. To mitigate against such extinction scenarios, we advocate the creation of a Chinese national agency and repository of environmental information focused on public awareness and education, the implementation of targeted programs of habitat restoration designed to return impacted forests to a more natural state especially within and at the boundaries of nature reserves, the establishment of additional protect areas, and the construction of a latticework of corridors connecting isolated primate subpopulations. This comprehensive approach offers the most effective way to protect China’s animal and plant biodiversity, including its endangered primate populations.

Evolution of Acidic Mammalian Chitinase Genes (CHIA) Is Related to Body Mass and Insectivory in Primates.

Insects are an important food resource for many primates, but the chitinous exoskeletons of arthropods have long been considered to be indigestible by the digestive enzymes of most mammals. However, recently mice and insectivorous bats were found to produce the enzyme acidic mammalian chitinase (AMCase) to digest insect exoskeletons. Here, we report on the gene CHIA and its paralogs, which encode AMCase, in a comparative sample of nonhuman primates. Our results show that early primates likely had three CHIA genes, suggesting that insects were an important component of the ancestral primate diet. With some exceptions, most extant primate species retain only one functional CHIA paralog. The exceptions include two colobine species, in which all CHIA genes have premature stop codons, and several New World monkey species that retain two functional genes. The most insectivorous species in our sample also have the largest number of functional CHIA genes. Tupaia chinensis and Otolemur garnettii retain three functional CHIA paralogs, whereas Tarsius syrichta has a total of five, two of which may be duplications specific to the tarsier lineage. Selection analyses indicate that CHIA genes are under more intense selection in species with higher insect consumption, as well as in smaller-bodied species (<500 g), providing molecular support for Kay's Threshold, a well-established component of primatological theory which proposes that only small primates can be primarily insectivorous. These findings suggest that primates, like mice and insectivorous bats, may use the enzyme AMCase to digest the chitin in insect exoskeletons, providing potentially significant nutritional benefits.

Functional analyses of the primate upper cervical vertebral column

Recent work has highlighted functional correlations between direct measures of head and neck posture and primate cervical bony morphology. Primates with more horizontal necks exhibit middle and lower cervical vertebral features that indicate increased mechanical advantage for deep nuchal musculature and mechanisms for column curvature formation and maintenance. How features of the C1 and C2 reflect quantified measures of posture have yet to be examined. This study incorporates bony morphology from the upper cervical levels from 20 extant primate species in order to investigate further how posture correlates with cervical vertebrae morphology. Results from phylogenetic generalized least-squares analyses indicate that few vertebral features exhibit a significant relationship with posture when accounting for differences in size. When size-adjusted traits were correlated with posture, vertebral variation had a stronger relationship with neck posture than head posture variables. Two C1 traits—relative posterior arch length and superior facet curvature—were correlated with neck posture variables. Relative posterior arch length exhibits a positive relationship with neck posture, while superior articular facet curvature demonstrates a negative relationship, such that as the neck becomes more horizontal, the greater the facet curvature. Four C2 features were also correlated with neck posture: relative pedicle and lamina lengths, relative superior facet orientation, and dens orientation. Relative pedicle and lamina lengths become craniocaudally longer as the neck becomes more horizontal. Relative C2 superior facet orientation and dens orientation exhibit negative correlations with posture, such that as the neck becomes more horizontal, the superior facet becomes more caudally inclined and the dens more dorsally inclined. These results produce a similar functional signal observed in the middle and lower cervical spine. Modeling the cervical vertebrae of more pronograde taxa within a sigmoidal spinal column model is further discussed and may prove useful in refining and testing future hypotheses of primate cervical mechanics.

Capuchin monkeys can make and use stone tools.

Scientists hoping to elucidate the origin of human stone tool manufacture and use have looked to extant primate species for possible clues. Although some skepticism has been raised, there is clear evidence that today's capuchin monkeys can make and use stone tools.