Ecological factors are likely drivers of eye shape and colour pattern variations across anthropoid primates.

External eye appearance across primate species is diverse in shape and colouration, yet we still lack an explanation for the drivers of such diversity. Here we quantify substantial interspecific variation in eye shape and colouration across 77 primate species representing all extant genera of anthropoid primates. We reassess a series of hypotheses aiming to explain ocular variation in horizontal elongation and in colouration across species. Heavier body weight and terrestrial locomotion are associated with elongated eye outlines. Species living closer to the equator present more pigmented conjunctivae, suggesting photoprotective functions. Irises become bluer in species living further away from the equator, adding to existing literature supporting a circadian clock function for bluer irises. These results shift the current focus from communicative, to ecological factors in driving variation in external eye appearance in anthropoid primates. They also highlight the possibility that similar ecological factors contributed to selection for blue eyes in ancestral human populations living in northern latitudes.

Morphological modularity in the anthropoid axial skeleton.

Previous research has found that hominoids have stronger modularity between limb elements than other anthropoids, suggesting that there is less constraint on morphological diversification (e.g., limb proportions) in hominoids in terms of evolutionary independence. However, degrees of modularity in the axial skeleton have not been investigated across a broad range of anthropoid taxa. Thus, it is unknown whether hominoids also have stronger modularity in the axial skeleton than other anthropoids, which has implications for the evolution of diverse torso morphologies in Miocene apes as well as the evolution of novel characteristics in the skull and vertebrae of fossil hominins. In this study, 12 anthropoid genera were sampled to examine degrees of modularity between axial skeletal elements (i.e., cranium, mandible, vertebrae, and sacrum). Covariance ratio coefficients were calculated using variance/covariance matrices of interlandmark distances for each axial skeletal element to evaluate degrees of modularity. The results showed that Alouatta, Hylobates, Gorilla, Pan, and Homo showed generally stronger modularity than other anthropoid taxa when considering all axial skeletal elements. When only considering the vertebral elements (i.e., vertebrae and sacrum), Alouatta, Hylobates, Gorilla, and Pan showed generally stronger modularity than other anthropoid taxa. Humans showed stronger modularity between the skull and vertebrae than other hominoids. Thus, the evolution of novel characteristics in the skull and vertebral column may have been less constrained in fossil hominins due to the dissociation of trait covariation between axial skeletal elements in hominoid ancestors, thus fostering more evolutionary independence between the skull and vertebral column.

Homo naledi pollical metacarpal shaft morphology is distinctive and intermediate between that of australopiths and other members of the genus Homo.

Homo naledi fossils from the Rising Star cave system provide important insights into the diversity of hand morphology within the genus Homo. Notably, the pollical (thumb) metacarpal (Mc1) displays an unusual suite of characteristics including a median longitudinal crest, a narrow proximal base, and broad flaring intrinsic muscle flanges. The present study evaluates the affinities of H. naledi Mc1 morphology via 3D geometric morphometric analysis of shaft shape using a broader comparative sample (n = 337) of fossil hominins, recent humans, apes, and cercopithecoid monkeys than in prior work. Results confirm that the H. naledi Mc1 is distinctive from most other hominins in being narrow at the proximal end but surmounted by flaring muscle flanges distally. Only StW 418 (Australopithecus cf. africanus) is similar in these aspects of shape. The gracile proximal shaft is most similar to cercopithecoids, Pan, Pongo, Australopithecus afarensis, and Australopithecus sediba, suggesting that H. naledi retains the condition primitive for the genus Homo. In contrast, Neandertal Mc1s are characterized by wide proximal bases and shafts, pinched midshafts, and broad distal flanges, while those of recent humans generally have straight shafts, less robust muscle flanges, and wide proximal shafts/bases. Although uncertainties remain regarding character polarity, the morphology of the H. naledi thumb might be interpreted as a retained intermediate state in a transformation series between the overall gracility of the shaft and the robust shafts of later hominins. Such a model suggests that the addition of broad medial and lateral muscle flanges to a primitively slender shaft was the first modification in transforming the Mc1 into the overall more robust structure exhibited by other Homo taxa including Neandertals and recent Homo sapiens in whose shared lineage the bases and proximal shafts became expanded, possibly as an adaptation to the repeated recruitment of powerful intrinsic pollical muscles.

A new parapithecine (Primates: Anthropoidea) from the early Oligocene of Libya supports parallel evolution of large body size among parapithecids.

Parapithecines are an extinct subfamily of stem anthropoid primates previously known only from the Jebel Qatrani Formation in Egypt. Here, we describe isolated teeth pertaining to Simonsius harujensis sp. nov., a relatively small-bodied parapithecine from strata near Zallah Oasis in the Sirt Basin of central Libya that is estimated to date to ∼31 Ma on the basis of mammalian biostratigraphy. The dental morphology of S. harujensis sp. nov. is generally intermediate between that of the closely related parapithecines Parapithecus fraasi and Simonsius grangeri, highlighting some of the anatomical features distinguishing the latter taxa and providing further support for their generic separation. A phylogenetic analysis using parsimony methods was performed on a character-taxon matrix incorporating data from the new Libyan parapithecine, virtually all other parapithecids and the proteopithecid Proteopithecus sylviae. Results of this analysis suggest that parapithecids comprise a basal clade consisting of three species of Biretia and a more derived clade including Parapithecinae (Parapithecus and Simonsius) and Qatraniinae (Qatrania, Ucayalipithecus, and Apidium). Body mass estimates for parapithecids were calculated on the basis of regression equations generated to predict body mass from the occlusal area of upper and lower cheek teeth in extant anthropoids. The relatively small body mass of S. harujensis sp. nov. and its reconstructed phylogenetic position as the sister group of S. grangeri, which is the largest known parapithecid, support the convergent acquisition of body mass larger than 500 g among multiple clades of early Oligocene African anthropoids. The new Libyan parapithecine augments previously reported evidence supporting a substantial degree of faunal provincialism across northern Africa/Arabia during the early Oligocene.

Mechanics of heel-strike plantigrady in African apes.

The initiation of a walking step with a heel strike is a defining characteristic of humans and great apes but is rarely found in other mammals. Despite the considerable importance of heel strike to an understanding of human locomotor evolution, no one has explicitly tested the fundamental mechanical question of why great apes use a heel strike. In this report, we test two hypotheses (1) that heel strike is a function of hip protraction and/or knee extension and (2) that short-legged apes with a midfoot that dorsiflexes at heel lift and long digits for whom digitigrady is not an option use heel-strike plantigrady. This strategy increases hip translation while potentially moderating the cost of redirecting the center of mass ('collisional costs') during stance via rollover along the full foot from the heel to toes. We quantified hind limb kinematics and relative hip translation in ten species of primates, including lemurs, terrestrial and arboreal monkeys, chimpanzees, and gorillas. Chimpanzees and gorillas walked with relatively extended knees but only with moderately protracted hips or hind limbs, partially rejecting the first hypothesis. Nonetheless, chimpanzees attained relative hip translations comparable with those of digitigrade primates. Heel-strike plantigrady may be a natural result of a need for increased hip translations when forelimbs are relatively long and digitigrady is morphologically restricted. In addition, foot rollover from the heel to toe in large, short-legged apes may reduce energetic costs of redirecting the center of mass at the step-to-step transition as it appears to do in humans. Heel strike appears to have been an important mechanism for increasing hip translation, and possibly reducing energetic costs, in early hominins and was fundamental to the evolution of the modern human foot and human bipedalism.

Variation of bony labyrinthine morphology in Mio-Plio-Pleistocene and modern anthropoids.

OBJECTIVES: The bony labyrinth of the inner ear has special relevance when tracking phenotypic evolution because it is often well preserved in fossil and modern primates. Here we track the evolution of the bony labyrinth of anthropoid primates during the Mio-Plio-Pleistocene-the time period that gave rise to the extant great apes and humans. MATERIALS AND METHODS: We use geometric morphometrics to analyze labyrinthine morphology in a wide range of extant and fossil anthropoids, including New World and Old World monkeys, apes, and humans; fossil taxa are represented by Aegyptopithecus, Microcolobus, Epipliopithecus, Nacholapithecus, Oreopithecus, Ardipithecus, Australopithecus, and Homo. RESULTS: Our results show that the morphology of the anthropoid bony labyrinth conveys a statistically significant phylogenetic signal especially at the family level. The bony labyrinthine morphology of anthropoids is also in part associated with size, but does not cluster by locomotor adaptations. The Miocene apes examined here, regardless of inferred locomotor behaviors, show labyrinthine morphologies distinct from modern great apes. DISCUSSION: Our results suggest that labyrinthine variation contains mixed signals and alternative explanations need to be explored, such as random genetic drift and neutral phenotypic evolution, as well as developmental constraints. The observed pattern in fossil and extant hominoids also suggests that an additional factor, for example, prenatal brain development, could have potentially had a larger role in the evolutionary modification of the bony labyrinth than hitherto recognized.

Is the middle cranial fossa a reliable predictor of temporal lobe volume in extant and fossil anthropoids?

OBJECTIVES: We investigate the suitability of middle cranial fossa (MCF) size as a proxy for temporal lobe volume (TLV), examining the strength of the association between TLV and MCF metrics and assess the reliability predicting TLV in fossil anthropoids. The temporal lobe of the primate brain is a multimodal association cortex involved in long-term memory, auditory, and visual processing with unique specializations in modern humans for language comprehension. The MCF is the bony counterpart for the temporal lobe providing inferences for fossil hominin temporal lobe evolution. We now investigate whether the MCF is a suitable proxy for the temporal lobe. METHODS: A sample of 23 anthropoid species (n = 232, including 13 fossil species) from computed tomography (CT) scans of ex vivo crania and magnetic resonance imaging (MRI) of the in vivo brain were generated into three-dimensional (3D) virtual models. Seven linear metrics were digitally measured on the right MCF with right TLV calculated from in vivo MRI. RESULTS: Regression analyses produced statistically significant correlations between TLV and all MCF metrics (r ≥ 0.85; p ≤ 0.0009) with TLV predictions within ±1 standard error and three MCF metrics (posterior-width, mid-length, and mid-width) the most reliable predictors of TLV with only one metric weakly associated with TLV. DISCUSSION: These findings indicate a strong association between the MCF and TLV, provide reliable predictors of fossil TLV that were previously unattainable, allow the inclusion of fragmentary fossil material, and enable inferences into the emergence of modern human temporal lobe morphology.

New Eocene primate from Myanmar shares dental characters with African Eocene crown anthropoids.

Recent discoveries of older and phylogenetically more primitive basal anthropoids in China and Myanmar, the eosimiiforms, support the hypothesis that Asia was the place of origins of anthropoids, rather than Africa. Similar taxa of eosimiiforms have been discovered in the late middle Eocene of Myanmar and North Africa, reflecting a colonization event that occurred during the middle Eocene. However, these eosimiiforms were probably not the closest ancestors of the African crown anthropoids. Here we describe a new primate from the middle Eocene of Myanmar that documents a new clade of Asian anthropoids. It possesses several dental characters found only among the African crown anthropoids and their nearest relatives, indicating that several of these characters have appeared within Asian clades before being recorded in Africa. This reinforces the hypothesis that the African colonization of anthropoids was the result of several dispersal events, and that it involved more derived taxa than eosimiiforms.

Cranial endocast of a stem platyrrhine primate and ancestral brain conditions in anthropoids.

Understanding of ancestral conditions for anthropoids has been hampered by the paucity of well-preserved early fossils. Here, we provide an unprecedented view of the cerebral morphology of the 20-million-year-old Chilecebus carrascoensis, the best-preserved early diverging platyrrhine known, obtained via high-resolution CT scanning and 3D digital reconstruction. These analyses are crucial for reconstructing ancestral brain conditions in platyrrhines and anthropoids given the early diverging position of Chilecebus. Although small, the brain of Chilecebus is not lissencephalic and presents at least seven pairs of sulci on its endocast. Comparisons of Chilecebus and other basal anthropoids indicate that the major brain subdivisions of these early anthropoids exhibit no consistent scaling pattern relative to the overall brain size. Many gross cerebral features appear to have transformed in a mosaic fashion and probably have originated in platyrrhine and catarrhine anthropoids independently, involving multiple, independent instances of size increase, as well as some secondary decreases.

Ontogeny of Orbit Orientation in Primates.

Orbit orientation in primates has been linked to adaptive factors related to activity pattern and size-related variation in structural influences on orbit position. Although differences in circumorbital form between anthropoids and strepsirrhines appear to be related to interspecific disparities in levels of orbital convergence and orbital frontation, there is considerable overlap in convergence between suborders. Unfortunately, putative links between convergence and frontation across primates, and consequent arguments about primate and anthropoid origins, are likely to be influenced by allometry, the size range of a respective sample, and adaptive influences on encephalization and activity patterns. Such a multifarious system is less amenable to interspecific treatment across higher-level clades. An ontogenetic perspective is one way to evaluate transformations from one character state to another, especially as they pertain to allometric effects on phenotypic variation. We characterized the ontogeny of orbital convergence and frontation in 13 anthropoid and strepsirrhine species. In each suborder, correlation and regression analyses were used to test hypotheses regarding the allometric bases of variation in orbital orientation. Growth trajectories were analyzed intra- and inter-specifically. Frontation decreased postnatally in all taxa due to the negative scaling of brain vs. skull size. Further, interspecific variation in relative levels of frontation was linked to corresponding ontogenetic transpositions in encephalization that differed within both suborders. In strepsirrhines, postnatal increases in convergence were largely due to the negative allometry of orbit vs. skull size. In contrast, convergence in anthropoids varied little during growth, being unrelated to ontogenetic variation in either relative orbit or interorbit size. Anat Rec, 302:2093-2104, 2019. © 2019 American Association for Anatomy.

Middle ear pneumatization in nonhuman primates: A comparative analysis.

OBJECTIVES: We test the effects of body mass and phylogeny on middle ear cavity pneumatization, and the role of pneumatization in hearing function, spanning the anatomical, ecological, and behavioral diversity of nonhuman primates. MATERIALS AND METHODS: All cavities were segmented in middle ear scans of 96 specimens, from 12 strepsirrhine and 15 haplorhine extant species. We measured the tympanic cavity (TC) separately, and all other middle ear spaces together (MES), calculating the degree of pneumatization with the surface area-to-volume ratio. We tested body mass effect with linear regression; we evaluated the phylogenetic signal and selection patterns, using a Kappa statistic test, and Ornstein-Uhlenbeck models (OU). We investigated the link between pneumatization and hearing sensitivity using phylogenetic regression. RESULTS: Testing body mass reveals an allometric pattern for both TC and MES dimensions. Degree of pneumatization in MES is dependent on body mass in haplorhines: larger animals have more pneumatized MES. Differences at various taxonomic ranks were observed for MES, while no phylogenetic influence was observed for TC. Infraorder selection patterns are different. Auditory performance is significantly related to degree of pneumatization, indicating that a pneumatized middle ear is associated with better perception of low frequencies. DISCUSSION: Pneumatization in MES is under differential selective pressure, indicating several optima for this trait. Pneumatization in MES probably modifies hearing sensitivity through pressure regulation mechanisms, auditory bulla size reduction, and frequency modulation. This could explain strepsirrhine adaptation to high-frequency perception, while haplorhine auditory perception is adapted to a broader sound range, including high and low frequencies.

New middle Eocene omomyines (Primates, Haplorhini) from San Diego County, California.

The Friars Formation of San Diego County, California, has yielded a middle Eocene mammalian fauna from the early part of the Uintan North American Land Mammal Age. Prior research on the primate fauna from the Friars Formation provides evidence of one notharctine and multiple omomyine species, but many specimens collected since the early 1980s remain unstudied. Here we describe three new omomyine genera from the Friars Formation. These new taxa range in estimated body mass from about 119 g to 757 g, and substantially expand the diversity of middle Eocene omomyoids known from Southern California. Resolution of the phylogenetic relationships of the new Friars Formation omomyines is complicated by the fact that different character-taxon matrices and tree building methods produce different results. Nevertheless, all preliminary phylogenetic analyses are congruent in recovering a close relationship between the three new genera and the omomyines Macrotarsius, Omomys, Ourayia, and Utahia. Prior research has documented a shift in omomyoid diversity in North America from the anantomophine-rich Bridgerian to the omomyine-rich Uintan. Our description of three new Uintan omomyine taxa from the Friars Formation further emphasizes these opposite trends in anaptomorphine and omomyine species richness during the middle Eocene. All three of the new taxa are currently known from only the Friars Formation in San Diego County, California. Four of the previously known omomyoid genera from Southern California (Dyseolemur, Chumashius, Yaquius, and Stockia) are also endemic to the region, further highlighting the provincial character of primate faunas in Utah, Southern California, and West Texas during the Uintan.

A probabilistic, distributed, recursive mechanism for decision-making in the brain.

Decision formation recruits many brain regions, but the procedure they jointly execute is unknown. Here we characterize its essential composition, using as a framework a novel recursive Bayesian algorithm that makes decisions based on spike-trains with the statistics of those in sensory cortex (MT). Using it to simulate the random-dot-motion task, we demonstrate it quantitatively replicates the choice behaviour of monkeys, whilst predicting losses of otherwise usable information from MT. Its architecture maps to the recurrent cortico-basal-ganglia-thalamo-cortical loops, whose components are all implicated in decision-making. We show that the dynamics of its mapped computations match those of neural activity in the sensorimotor cortex and striatum during decisions, and forecast those of basal ganglia output and thalamus. This also predicts which aspects of neural dynamics are and are not part of inference. Our single-equation algorithm is probabilistic, distributed, recursive, and parallel. Its success at capturing anatomy, behaviour, and electrophysiology suggests that the mechanism implemented by the brain has these same characteristics.

Three-dimensional anatomy of the anthropoid bony pelvis.

OBJECTIVES: Pelvic form is hypothesized to reflect locomotor adaptation in anthropoids. Most observed variation is found in the ilium, which traditionally is thought to reflect thoracic and shoulder morphology. This article examines the articulated bony pelvis of anthropoids in three dimensions (3D) to test hypothesized variation in pelvic anatomy related to overall torso form. MATERIALS AND METHODS: Sixty landmarks were collected on articulated pelves from 240 anthropoid individuals. Landmark data were subjected to a Generalized Procrustes Analysis. Principal Components Analysis was used to identify trends among taxa. Linear metrics were extracted, and bivariate allometric analysis was used to compare intergroup differences and scaling trends of specific dimensions. RESULTS: The combination of 3D and bivariate allometric analysis demonstrates a complex pattern of locomotor/phylogenetic and allometric influences on pelvic morphology. Apes have relatively narrower dorsal interiliac spacing than do most monkeys, with relatively smaller spinal muscle attachment areas but only minimally wider ventral bi-iliac breadths. Hylobatids and atelids have a relatively more cranial position of their sacra than do other taxa, and hylobatids and cercopithecids relatively more retroflexed ischia. Within groups, the three pelvic joints (lumbosacral, sacroiliac, and hip) become relatively closer together with increasing body size. CONCLUSIONS: A three-dimensional consideration of the articulated pelvis in anthropoids reveals determinants of pelvic variation not previously appreciated by studies of isolated hipbones. This study provides no support for the hypothesis that the ape pelvis is mediolaterally broader than that of monkeys in relative terms, as would be expected if iliac shape is related to hypothesized differences in thoracic breadth and shoulder orientation. Instead, apes, especially great apes, have relatively narrow sacra and longer lower pelves, related to their shorter, stiffer lumbar spines and torsos. This difference, coupled with strong positive allometry of iliac breadth and negative allometry of key pelvic lengths, along with some variation in ischial morphology in certain taxa, explains much of the variation in pelvic form among anthropoid primates.

Kinematics of the anthropoid os centrale and the functional consequences of scaphoid-centrale fusion in African apes and hominins.

In most primates, the os centrale is interposed between the scaphoid, trapezoid, trapezium, and head of the capitate, thus constituting a component of the wrist's midcarpal complex. Scaphoid-centrale fusion is among the clearest morphological synapomorphies of African apes and hominins. Although it might facilitate knuckle-walking by increasing the rigidity and stability of the radial side of the wrist, the exact functional significance of scaphoid-centrale fusion is unclear. If fusion acts to produce a more rigid radial wrist that stabilizes the hand and limits shearing stresses, then in taxa with a free centrale, it should anchor ligaments that check extension and radial deviation, but exhibit motion independent of the scaphoid. Moreover, because the centrale sits between the scaphoid and capitate (a major stabilizing articulation), scaphoid-centrale mobility should correlate with scaphocapitate mobility in extension and radial deviation. To test these hypotheses, the centrale's ligamentous binding was investigated via dissection in Pongo and Papio, and the kinematics of the centrale were quantified in a cadaveric sample of anthropoids (Pongo sp., Ateles geoffroyi, Colobus guereza, Macaca mulatta, and Papio anubis) using a computed-tomography-based method to track wrist-bone motion. Results indicate that the centrale rotates freely relative to the scaphoid in all taxa. However, centrale mobility is only correlated with scaphocapitate mobility during extension in Pongo-possibly due to differences in overall wrist configuration between apes and monkeys. If an extant ape-like wrist characterized early ancestors of African apes and hominins, then scaphoid-centrale fusion would have increased midcarpal rigidity in extension relative to the primitive condition. Although biomechanically consistent with a knuckle-walking hominin ancestor, this assumes that the trait evolved specifically for that biological role, which must be squared with contradictory interpretations of extant and fossil hominoid morphology. Regardless of its original adaptive significance, scaphoid-centrale fusion likely presented a constraint on early hominin midcarpal mobility.

Nasolacrimal anatomy and haplorhine origins.

Computed tomography X-ray imaging of the internal face in well-preserved primate fossil crania permits reconstruction of the nature of their nasal anatomy, including some soft-tissue features. These features are diagnostic of the primate suborder Haplorhini, and allow reevaluation of the phylogenetic status of several purported early members of the group. Here we examine the nasolacrimal morphology of a broad sample of extant primates, as well as a number of Paleogene fossils. The extant sample confirms the distinctiveness of the two suborders. Of the fossils studied, only Shoshonius cooperi from the late-early Eocene exhibits evidence of a haplorhine nose. This suggests that the haplorhine oronasal complex may have evolved before the postorbital septum, and strengthens the claim that Shoshonius is a close relative of tarsiers and anthropoids. These results indicate that Omomyiformes is not a monophyletic group, and that few of its members possessed the derived oronasal morphology that characterizes crown haplorhines.

Functional correlates of the position of the axis of rotation of the mandible during chewing in non-human primates.

The location of the axis of rotation (AoR) of the mandible was quantified using the helical axis (HA) in eight individuals from three species of non-human primates: Papio anubis, Cebus apella, and Macaca mulatta. These data were used to test three hypotheses regarding the functional significance of anteroposterior condylar translation - an AoR located inferior to the temporomandibular joint (TMJ) - during chewing: minimizing impingement of the gonial region on cervical soft tissue structures during jaw opening; avoiding stretching of the inferior alveolar neurovascular bundle (IANB); and increasing jaw-elevator muscle torques. The results reveal that the HA is located near the occlusal plane in Papio and Cebus, but closer to the condyle in Macaca; is located anteroinferior to the TMJ during both opening and closing in Papio, as well as during opening in Macaca and Cebus; and varies in its location during closing in Macaca and Cebus. The impingement hypothesis is not supported by interspecific variation in HA location: species with larger gonial angles like Cebus do not have more inferiorly located HAs than species with more obtuse mandibular angles like Papio. However, intraspecific variation provides some support for the impingement hypothesis. The HA seldom passes near or through the lingula, falsifying the hypothesis that its location is determined by the sphenomandibular ligament, and the magnitudes of strain associated with a HA at the TMJ would not be large enough to cause problematic stretching of the IANB. HA location does affect muscle moment arms about the TMJ, with implications for the torque generation capability of the jaw-elevator muscles. In Cebus, a HA farther away from the TMJ is associated with larger jaw-elevator muscle moment arms about the joint than if it were at the TMJ. The effects of HA location on muscle strain and muscle moment arms are largest at large gapes and smallest at low gapes, suggesting that if HA location is of functional significance for primate feeding system performance, it is more likely to be in relation to large gape feeding behaviors than chewing. Its presence in humans is most parsimoniously interpreted as a primitive retention from non-human primate ancestors and explanations for the presence of anteroposterior condylar translation in humans need not invoke either the uniqueness of human speech or upright posture.

Revisiting size and scaling in the anthropoid temporomandibular joint.

Masticatory morphology in primates is likely under strong selective pressure to maximize feeding efficiency while simultaneously minimizing the occurrence of injury or pathology. As a result, masticatory shape, including aspects of temporomandibular joint (TMJ) morphology, varies widely across primates in relation to feeding behavior and body size. This study examines patterns of allometry in the TMJ of anthropoid primates, with the specific goal of evaluating how allometric patterns may reflect variation in loading and/or range of motion at this joint. Phylogenetic reduced major axis regressions were employed to examine how specific aspects of TMJ morphology scale in relation to body mass and mandible length. Patterns of shape variation across the entire masticatory apparatus were examined by utilizing geometric morphometric techniques. Results reveal that most aspects of TMJ shape scale with either isometry or positive allometry relative to body mass and/or mandible length, though several departures from these patterns were observed. In particular, male cercopithecoids tend to show distinct scaling patterns in TMJ height above the occlusal plane and condylar area, likely reflecting known trade-offs between increased range of motion and force production in this clade, as has been linked to selection for increased male canine size. The geometric morphometric analyses indicate that craniofacial and masticatory shape are strongly allometric, but that glenoid shape variation is less consistently allometric. Notably, different patterns of allometric shape variation were observed in platyrrhines, cercopithecoids, and hominoids, perhaps related to different, and potentially competing, selective pressures in each of these clades.

Genetics of Cerebellar and Neocortical Expansion in Anthropoid Primates: A Comparative Approach.

What adaptive changes in brain structure and function underpin the evolution of increased cognitive performance in humans and our close relatives? Identifying the genetic basis of brain evolution has become a major tool in answering this question. Numerous cases of positive selection, altered gene expression or gene duplication have been identified that may contribute to the evolution of the neocortex, which is widely assumed to play a predominant role in cognitive evolution. However, the components of the neocortex co-evolve with other functionally interdependent regions of the brain, most notably in the cerebellum. The cerebellum is linked to a range of cognitive tasks and expanded rapidly during hominoid evolution. Here we present data that suggest that, across anthropoid primates, protein-coding genes with known roles in cerebellum development were just as likely to be targeted by selection as genes linked to cortical development. Indeed, based on currently available gene ontology data, protein-coding genes with known roles in cerebellum development are more likely to have evolved adaptively during hominoid evolution. This is consistent with phenotypic data suggesting an accelerated rate of cerebellar expansion in apes that is beyond that predicted from scaling with the neocortex in other primates. Finally, we present evidence that the strength of selection on specific genes is associated with variation in the volume of either the neocortex or the cerebellum, but not both. This result provides preliminary evidence that co-variation between these brain components during anthropoid evolution may be at least partly regulated by selection on independent loci, a conclusion that is consistent with recent intraspecific genetic analyses and a mosaic model of brain evolution that predicts adaptive evolution of brain structure.

New cranium of the endemic Caribbean platyrrhine, Antillothrix bernensis, from La Altagracia Province, Dominican Republic.

Recent paleontological collection in submerged caves in the eastern Dominican Republic has yielded new specimens of Antillothrix bernensis. Here we describe a complete cranium of an adult individual (MHD 20) and provide phenetic comparisons to other endemic Caribbean taxa and extant mainland platyrrhines using three-dimensional geometric morphometric methods (3DGM). Qualitative and quantitative comparisons support conclusions based on other recently described fossil material: Antillothrix has a dentition lacking clear dietary specialization, an elongated brain case with strong temporal lines, and a vertically oriented nuchal plane. MHD 20 shares a combination of traits with a previously published subadult specimen (MHD 01) including a deep depression at glabella, dorsoventrally elongated orbits, and a relatively large face. This shared morphology reinforces the taxonomic affinity of the two specimens, with differences between the two likely reflecting the younger ontogenetic age of MHD 01. Comparisons to the extant platyrrhines paint a complicated picture as the results of between-group principal components analyses (bgPCA) indicate that Antillothrix does not share a suite of morphological features exclusively with any one genus. Depending on which bgPC axes are visualized, and which subset of landmarks is included (i.e., only those describing the shape of the face/palate for inclusion of Xenothrix), MHD 20 is most similar in shape to the atelids, Alouatta, Lagothrix, and Brachyteles, or an otherwise "empty" region of shape space. It groups neither with Cebus nor Callicebus, two taxa that Antillothrix has been associated with in previous studies based on much less complete material. The Antillothrix cranium does not exhibit any of the derived characters classically used to diagnose or define any single clade; rather its morphology shares features with multiple platyrrhine groups. This is consistent with the interpretation that Antillothrix preserves a primitive morphology, which accords with the hypothesis positing an early arrival of platyrrhines in the Caribbean.