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For several decades, it was largely assumed that stone tool use and production were abilities limited to the genus Homo. However, growing palaeontological and archaeological evidence, comparative extant primate studies, as well as results from methodological advancements in biomechanics and morphological analyses, have been gradually accumulating and now provide strong support for more advanced manual manipulative abilities and tool-related behaviours in pre-Homo hominins than has been traditionally recognized. Here, I review the fossil evidence related to early hominin dexterity, including the recent discoveries of relatively complete early hominin hand skeletons, and new methodologies that are providing a more holistic interpretation of hand function, and insight into how our early ancestors may have balanced the functional requirements of both arboreal locomotion and tool-related behaviours. [ABSTRACT FROM AUTHOR]

Oldowan lithic assemblages are often portrayed as a product of the need to obtain sharp flakes for cutting into animal car cases. However, ethnographic and experimental research indicates that the optimal way to produce flakes for such butchering purposes is via bipolar reduction of small cryptocrystalline pebbles rather than from larger crystalline cores resembling choppers. Ethnographic observations of stone tool-using hunter-gatherers in environments comparable with early hominins indicate that most stone tools (particularly chopper forms and flake tools) were used for making simple shaft tools including spears, digging sticks and throwing sticks. These tools bear strong resemblances to Oldowan stone tools. Bipolar reduction for butchering probably preceded chopper-like core reduction and provides a key link between primate nut-cracking technologies and the emergence of more sophisticated lithic technologies leading to the Oldowan. [ABSTRACT FROM AUTHOR]

Recovering evidence of past human activities enables us to recreate behaviour where direct observations are missing. Here, we apply archaeological methods to further investigate cultural transmission processes in percussive tool use among neighbouring chimpanzee communities in the Taï National Park, Côte d'Ivoire, West Africa. Differences in the selection of nut-cracking tools between neighbouring groups are maintained over time, despite frequent female transfer, which leads to persistent cultural diversity between chimpanzee groups. Through the recovery of used tools in the suggested natal territory of immigrants, we have been able to reconstruct the tool material selection of females prior to migration. In combination with direct observations of tool selection of local residents and immigrants after migration, we uncovered temporal changes in tool selection for immigrating females. After controlling for ecological differences between territories of immigrants and residents our data suggest that immigrants abandoned their previous tool preference and adopted the pattern of their new community, despite previous personal proficiency of the same foraging task. Our study adds to the growing body of knowledge on the importance of conformist tendencies in animals. [ABSTRACT FROM AUTHOR]

Percussive technology is part of the behavioural suite of several fossil and living primates. Stone Age ancestors used lithic artefacts in pounding activities, which could have been most important in the earliest stages of stone working. This has relevant evolutionary implications, as other primates such as chimpanzees and some monkeys use stone hammer-and-anvil combinations to crack hard- shelled foodstuffs. Parallels between primate percussive technologies and early archaeological sites need to be further explored in order to assess the emergence of technological behaviour in our evolutionary line, and firmly establish bridges between Primatology and Archaeology. What are the anatomical, cognitive and ecological constraints of percussive technology? How common are percussive activities in the Stone Age and among living primates? What is their functional significance? How similar are archaeological percussive tools and those made by non-human primates? This issue of Phil. Trans. addresses some of these questions by presenting case studies with a wide chronological, geographical and disciplinary coverage. The studies presented here cover studies of Brazilian capuchins, captive chimpanzees and chimpanzees in the wild, research on the use of percussive technology among modern humans and recent hunter-gatherers in Australia, the Near East and Europe, and archaeological examples of this behaviour from a million years ago to the Holocene. In summary, the breadth and depth of research compiled here should make this issue of Philosophical Transactions of the Royal Society B, a landmark step forward towards a better understanding of percussive technology, a unique behaviour shared by some modern and fossil primates. [ABSTRACT FROM AUTHOR]

Various authors have suggested similarities between tool use in early hominins and chimpanzees. This has been particularly evident in studies of nut-cracking which is considered to be the most complex skill exhibited by wild apes, and has also been interpreted as a precursor of more complex stone-flaking abilities. It has been argued that there is no major qualitative difference between what the chimpanzee does when he cracks a nut and what early hominins did when they detached a flake from a core. In this paper, similarities and differences between skills involved in stone-flaking and nut-cracking are explored through an experimental protocol with human subjects performing both tasks. We suggest that a 'functional' approach to percussive action, based on the distinction between functional parameters that characterize each task and parameters that characterize the agent's actions and movements, is a fruitful method for understanding those constraints which need to be mastered to perform each task successfully, and subsequently, the nature of skill involved in both tasks. [ABSTRACT FROM AUTHOR]

Percussive tool use holds special interest for scientists concerned with human origins. We summarize the findings from two field sites, Taï and Fazenda Boa Vista, where percussive tool use by chimpanzees and bearded capuchins, respectively, has been extensively investigated. We describe the ecological settings in which nut-cracking occurs and focus on four aspects of nut-cracking that have important cognitive implications, namely selection of tools, tool transport, tool modification and modulation of actions to reach the goal of cracking the nut. We comment on similarities and differences in behaviour and consider whether the observed differences reflect ecological, morphological, social and/or cognitive factors. Both species are sensitive to physical properties of tools, adjust their selection of hammers conditionally to the resistance of the nuts and to transport distance, and modulate the energy of their strikes under some conditions. However, chimpanzees transport hammers more frequently and for longer distances, take into account a higher number of combinations of variables and occasionally intentionally modify tools. A parsimonious interpretation of our findings is that morphological, ecological and social factors account for the observed differences. Confirmation of plausible cognitive differences in nut-cracking requires data not yet available. [ABSTRACT FROM AUTHOR]

The skill of object manipulation is a common feature of primates including humans, although there are species-typical patterns of manipulation. Object manipulation can be used as a comparative scale of cognitive development, focusing on its complexity. Nut cracking in chimpanzees has the highest hierarchical complexity of tool use reported in non-human primates. An analysis of the patterns of object manipulation in naive chimpanzees after nut-cracking demonstrations revealed the cause of difficulties in learning nut-cracking behaviour. Various types of behaviours exhibited within a nut-cracking context can be examined in terms of the application of problem-solving strategies, focusing on their basis in causal understanding or insightful intentionality. Captive chimpanzees also exhibit complex forms of combinatory manipulation, which is the precursor of tool use. A new notation system of object manipulation was invented to assess grammatical rules in manipulative actions. The notation system of action grammar enabled direct comparisons to be made between primates including humans in a variety of object-manipulation tasks, including percussive-tool use. [ABSTRACT FROM AUTHOR]

Pliopithecoids represent a monophyletic group of putative stem catarrhines whose evolutionary history is incompletely known. They have been recorded from Europe and Asia, between the late Early Miocene and the Late Miocene. Asian pliopithecoids are less well documented than their European counterparts, often being represented by a fragmentary fossil record. New discoveries are therefore critical to reconstruct the evolutionary history of the whole group. Here, we describe two isolated molars from Ban San Klang, a late Middle Miocene locality in northern Thailand, which confirms the presence of pliopithecoids in Southeast Asia. The lower molar had originally been described as being that of a dendropithecoid, but it was later recognized as pertaining to a pliopithecoid. The discovery, in the same locality, of an additional upper molar attributed to the same species confirms the pliopithecoid status of this taxon and highlights its distinctiveness with respect to other known Asian pliopithecoids. However, the mosaic of primitive and autapomorphic features characterizing this Thai fossil, as well as its limited anatomical representation, preclude us from assigning it to either of the known pliopithecid subfamilies. Nevertheless, it represents the only pliopithecoid in Southeast Asia and displays a mosaic of unique characters which emphasizes the peculiarity of that province, as suggested previously with respect to its hominoid primate. [ABSTRACT FROM AUTHOR]

A variety of lines of evidence support the idea that neutral evolutionary processes (genetic drift, mutation) have been important in generating cranial differences between Neandertals and modern humans. But how do Neandertals and modern humans compare with other species? And how do these comparisons illuminate the evolutionary processes underlying cranial diversification? To address these questions, we used 27 standard cranial measurements collected on 2524 recent modern humans, 20 Neandertals and 237 common chimpanzees to estimate split times between Neandertals and modern humans, and between Pan troglodytes verus and two other subspecies of common chimpanzee. Consistent with a neutral divergence, the Neandertal versus modern human split-time estimates based on cranial measurements are similar to those based on DNA sequences. By contrast, the common chimpanzee cranial estimates are much lower than DNA-sequence estimates. Apparently, cranial evolution has been unconstrained in Neandertals and modern humans compared with common chimpanzees. Based on these and additional analyses, it appears that cranial differentiation in common chimpanzees has been restricted by stabilizing natural selection. Alternatively, this restriction could be due to genetic and/or developmental constraints on the amount of within-group variance (relative to effective population size) available for genetic drift to act on. [ABSTRACT FROM AUTHOR]

Eating is a risky affair. All animals have to offset risks of feeding such as exposure to plant toxins, increased vulnerability to predation, or conspecific aggression with a food's energetic and nutritional return. What, when, and where an individual eats can impact fitness and, ultimately, species-level adaptations. Here, we explore the variables that influence primate feeding preference: food availability, chemical defense, and nutrient content. We present information demonstrating that consumers manipulate nutrient and energy intake, indicating that what may be a less-than-optimal food for one state of an animal's phenotype may not be for another. This evidence suggests that factors previously assumed to be constraints in Optimal Foraging Theory, Functional Response, and-recently-Fallback Food feeding models would be better categorized as variables. We conclude that 'fallback' is not an intrinsic state of the food or the consumer and that this conclusion complicates the application of this concept to morphological features in the fossil record. [ABSTRACT FROM AUTHOR]

ABSTRACT Objectives Little is known about the absolute pool of energy primates use in the wild and how this pool varies seasonally. It is also not fully understood why some species facing unpredictable changes in food supply and risks of energy imbalance show high breeding seasonality. We examined these issues in a folivorous gregarious lemur ( Propithecus coronatus) in a dry forest of Madagascar. Methods We assessed energy input and activity budget in males and females from three groups during dry and wet months. Diet composition and daily food intake were determined using focal animal sampling and continuous recording of mouthfuls. Locomotion types and daily distances traveled were recorded. Time budget was determined using scan sampling, supplemented with a preliminary analysis of the energy budget using the factorial method. Results The macronutrient composition of the diet did not change markedly between seasons but a large increase in energy input arose during the early wet season. Activity was highest at this period but entailed only moderate increase in energy expenditure. Females did not ingest more energy than males despite female dominance. Both sexes exhibited a thrifty energy strategy and, possibly, fat storage. Conclusions Seasonal increase of daily energy input was likely related to male-male mating competition and females' need to reach physical condition prior to the dry season sufficient for reproducing. Sifakas may time their reproductive cycle according to the seasonal variation in day-length because recurrent patterns of low (dry season) versus high (wet season) food production occur despite years of food shortage. Am J Phys Anthropol, 2015. © 2015 Wiley Periodicals, Inc. Am J Phys Anthropol 158:269-278, 2015. © 2015 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

ABSTRACT Objective The African papionin primates commonly known as mangabeys form a diphyletic group with white-eyelid mangabeys ( Cercocebus) being most closely related to drills and mandrills ( Mandrillus). However, the phylogenetic relationships among members of the Cercocebus-Mandrillus clade have not been investigated in detail, particularly from a morphological perspective. Early studies of white-eyelid mangabeys considered C. agilis to best represent the ancestral lineage and C. torquatus as the most derived species, the result of multiple biogeographic dispersal events. More recently, a sister-clade relationship between Mandrillus and either C. chrysogaster or C. torquatus has been proposed. Materials and Methods Here we present the results of phylogenetic analyses based on 206 craniodental characters (103 representing males and females separately) of four species of Cercocebus and both species of Mandrillus. Results When all species of the Cercocebus-Mandrillus clade are analyzed without molecular constraints, results suggest that C. torquatus may indeed be more closely related to mandrills and drills than to other Cercocebus taxa. However, this topology lacks strong statistical support and may be attributable to incomplete lineage sorting and/or reticulation. When we employ a molecular backbone to constrain Cercocebus and Mandrillus monophyly, C. torquatus appears most basal, while C. agilis and C. chrysogaster form a derived clade. Discussion The molecular backbone view is also congruent with recent genetic analyses and assessments based on the fossil record, but awaits confirmation with additional data. This phylogeny suggests that Cercocebus and Cercocebus + Mandrillus arose in western equatorial Africa with subsequent dispersals westward, eastward, and possibly southward over the last > 3 million years. Am J Phys Anthropol 158:227-241, 2015. © 2015 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

Although some people can voluntarily move their ears, overt reflexive control of the pinnae has been lost during the course of primate evolution. Humans and apes do not move their ears to express emotion, they do not defensively retract them when startled, and they do not point them at novel, salient, or task-relevant stimuli. Nevertheless, it is the thesis of this review that neural circuits for pinna orienting have survived in a purely vestigial state for over 25 million years. There are three lines of evidence: (1) Shifting the eyes hard to one side is accompanied by electromyographic (EMG) activity in certain ear muscles and by a barely visible (2-3 mm) curling of the dorsal edge of the pinna. (2) The capture of attention by a novel, unexpected sound emanating from behind and to one side has been found to trigger a weak EMG response in the muscle behind the corresponding ear. (3) Reflexive EMG bursts recorded during a selective attention task suggested that subjects were unconsciously attempting to orient their ears toward the relevant sounds. In addition to pinna orienting, the possibility that pinna startle might have survived in a vestigial state is also considered. It is suggested that the postauricular reflex to sudden, intense sounds constitutes a vestigial startle response, but that the reflex arc is dominated by a pathway that bypasses the main organizing center for startle. [ABSTRACT FROM AUTHOR]

ABSTRACT The need to maintain stability on narrow branches is often presented as a major selective force shaping primate morphology, with adaptations to facilitate grasping receiving particular attention. The functional importance of a long and mobile tail for maintaining arboreal stability has been comparatively understudied. Tails can facilitate arboreal balance by acting as either static counterbalances or dynamic inertial appendages able to modulate whole-body angular momentum. We investigate associations between tail use and inferred grasping ability in two closely related cebid platyrrhines-cotton-top tamarins ( Saguinus oedipus) and black-capped squirrel monkeys ( Saimiri boliviensis). Using high-speed videography of captive monkeys moving on 3.2 cm diameter poles, we specifically test the hypothesis that squirrel monkeys (characterized by grasping extremities with long digits) will be less dependent on the tail for balance than tamarins (characterized by claw-like nails, short digits, and a reduced hallux). Tamarins have relatively longer tails than squirrel monkeys, move their tails through greater angular amplitudes, at higher angular velocities, and with greater angular accelerations, suggesting dynamic use of tail to regulate whole-body angular momentum. By contrast, squirrel monkeys generally hold their tails in a comparatively stationary posture and at more depressed angles, suggesting a static counterbalancing mechanism. This study, the first empirical test of functional tradeoffs between grasping ability and tail use in arboreal primates, suggests a critical role for the tail in maintaining stability during arboreal quadrupedalism. Our findings have the potential to inform our functional understanding of tail loss during primate evolution. J. Exp. Zool. 323A: 556-566, 2015. © 2015 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

Climatic niches have increasingly become a nexus in our understanding of a variety of ecological and evolutionary phenomena, from species distributions to latitudinal diversity gradients. Despite the increasing availability of comprehensive datasets on species ranges, phylogenetic histories, and georeferenced environmental conditions, studies on the evolution of climate niches have only begun to understand how niches evolve over evolutionary timescales. Here, using primates as a model system, we integrate recently developed phylogenetic comparative methods, species distribution patterns, and climatic data to explore primate climatic niche evolution, both among clades and over time. In general, we found that simple, constant-rate models provide a poor representation of how climatic niches evolve. For instance, there have been shifts in the rate of climatic niche evolution in several independent clades, particularly in response to the increasingly cooler climates of the past 10 My. Interestingly, rate accelerations greatly outnumbered rate decelerations. These results highlight the importance of considering more realistic evolutionary models that allow for the detection of heterogeneity in the tempo and mode of climatic niche evolution, as well as to infer possible constraining factors for species distributions in geographical space. [ABSTRACT FROM AUTHOR]

Primates often make foraging selections that are not apparent. For example, they may eagerly consume a particular plant part and species in some instances, but reject it at other times. Blighia unijugata (Baker) fruit is one of the most frequently eaten foods of mangabeys (Lophocebus albigena) in Lwamunda Forest Reserve, Uganda; however, its use varies strikingly depending on the fruit's developmental stage. We conducted feeding observations to investigate the nutritional criteria that mangabeys may have used for the consumption of specific fruit parts by conducting analysis of fruit parts eaten and rejected at different developmental stages. When seeds had low condensed tannins, mangabeys ate them, but seeds were rejected when tannin levels increased. In the first five stages of fruit development, tannin levels in arils declined and the frequency of consumption of the aril increased. Although fat content in seeds increased with maturity, it did not appear to influence seed consumption, but fat content was related to frequency of consumption of the aril. Considering that primates are often making food selections among many species/part combinations, our results illustrate the value of using nutritional analyses to understand foraging decisions. Furthermore, they demonstrate how very specific trade-offs between consumption of critical nutrients and antifeedants can drive which foods and parts are eaten. [ABSTRACT FROM AUTHOR]

In most anthropoid primates, the maxillary canine, mandibular canine, and mesial mandibular premolar form a functional complex that hones the canines. Characters in functional complexes are predicted to covary genetically, which constrains their evolutionary independence. As a result of substantial changes to canine and honing premolar size and shape, hominins are characterized by the apomorphic loss of canine honing. In early hominins, changes in canine and ‘honing’ premolar size and shape appear to have been uncoordinated, which is unexpected if there is strong genetic covariation coupling these teeth. Using the pattern and magnitude of phenotypic dental size covariation in extant anthropoids, results of this study indicate that certain dimensions of the anthropoid honing complex are characterized by strong size covariation within species and that canine and honing premolar size have evolved in a coordinated manner in both males and females, which undermines arguments that the complex is selectively important only in males. Further, there is no evidence for negative or strong positive covariance between canine and either incisor or postcanine size. If patterns of phenotypic covariation reflect genetic covariation, this suggests that canine reduction was unlikely to have been a dependent change associated with the development of postcanine megadontia or incisor reduction. [ABSTRACT FROM AUTHOR]

Research on primate origins has revolved around arboreality and, more specifically, the adaptations that are linked to safe navigation in the fine-branch niche. To this end, extant non-primate mammals have been used as models to assess the significance of these adaptations. However, the size of these models is larger than that estimated for early primates. In contrast, the feathertail marsupial glider Acrobates pygmaeus , with a body mass of 12 g, a clawless opposable hallux, and terminal branch feeding habits appears more suited to modeling behavioral adaptations to the small branch milieu. Analysis of video recordings of 18 feathertail gliders walking on poles of variable diameter and inclination revealed that they preferentially used diagonal sequence gaits, fast velocities and low duty factors. Diagonal gaits did not correlate to duty factor, but increased as substrate size decreased, and from descending to ascending locomotion. Furthermore, the duty factor index increased in more diagonal gaits and ascending locomotion. Finally, velocities were lower on smaller substrates, and were mainly regulated by stride frequency and, to a lesser degree, stride length. Feathertail glider gaits displayed noteworthy behavioral convergences with primate quadrupedalism, but some of these results need additional investigation. Despite any discrepancies, these features appear to be favorable for quadrupedal progression on small branches, providing a selective advantage for navigating within a fine branch niche and highlighting the importance of small body size in early primate evolution. [ABSTRACT FROM AUTHOR]

Early primate evolution is connected to the efficient exploitation of the terminal branch habitat. Mammals that forage in this habitat constantly encounter flexible thin branches that bend under the weight of the animals and thus form steeply inclined and declined supports. This study was aimed to gain insight into how cotton-top tamarins - a previously proposed modern analogue for a hypothetical stage in early primate evolution with prehensile autopodia - meet the specific functional demands when navigating thin, branch-like supports of different orientation. X-ray motion analysis was combined with synchronous single limb substrate reaction force measurements to discern limb mechanical adjustments. Previously reported gait parameter adjustments were confirmed for moderate support orientations, but on the steepest inclines and declines kinematic discontinuities were observed. These are interpreted to emphasize the functional roles of the forelimbs (net-braking role) and hind limbs (net-propulsive role) already established for level and moderately inclined supports. Tensile forces were exerted by the forelimbs on the steepest inclines and by the hind limbs on the steepest declines (head-first descents). Even though non-specialized small mammals have also been shown to successfully negotiate similar supports, prehensility offers advantages for foraging on thin, steeply inclined and declined terminal branches. Thus, the evolution of prehensile autopodia in small early primates likely has enhanced the exploitation of the terminal branch habitat. [ABSTRACT FROM AUTHOR]

The evolution of primates involved increasing body size, brain size and presumably cognitive ability. Cognition is related to neural activity, metabolic rate and rate of blood flow to the cerebral cortex. These parameters are difficult to quantify in living animals. This study shows that it is possible to determine the rate of cortical brain perfusion from the size of the internal carotid artery foramina in skulls of certain mammals, including haplorrhine primates and diprotodont marsupials. We quantify combined blood flow rate in both internal carotid arteries as a proxy of brain metabolism in 34 species of haplorrhine primates (0.116-145 kg body mass) and compare it to the same analysis for 19 species of diprotodont marsupials (0.014-46 kg). Brain volume is related to body mass by essentially the same exponent of 0.70 in both groups. Flow rate increases with haplorrhine brain volume to the 0.95 power, which is significantly higher than the exponent (0.75) expected for most organs according to 'Kleiber's Law'. By comparison, the exponent is 0.73 in marsupials. Thus, the brain perfusion rate increases with body size and brain size much faster in primates than in marsupials. The trajectory of cerebral perfusion in primates is set by the phylogenetically older groups (New and Old World monkeys, lesser apes) and the phylogenetically younger groups (great apes, including humans) fall near the line, with the highest perfusion. This may be associated with disproportionate increases in cortical surface area and mental capacity in the highly social, larger primates. [ABSTRACT FROM AUTHOR]

Skeletal histology supports the hypothesis that primate life histories are regulated by a neuroendocrine rhythm, the Havers-Halberg Oscillation (HHO). Interestingly, subfossil lemurs are outliers in HHO scaling relationships that have been discovered for haplorhine primates and other mammals. We present new data to determine whether these species represent the general lemur or strepsirrhine condition and to inform models about neuroendocrine-mediated life history evolution. We gathered the largest sample to date of HHO data from histological sections of primate teeth (including the subfossil lemurs) to assess the relationship of these chronobiological measures with life history-related variables including body mass, brain size, age at first female reproduction, and activity level. For anthropoids, these variables show strong correlations with HHO conforming to predictions, though body mass and endocranial volume are strongly correlated with HHO periodicity in this group. However, lemurs (possibly excepting Daubentonia) do not follow this pattern and show markedly less variability in HHO periodicity and lower correlation coefficients and slopes. Moreover, body mass is uncorrelated, and brain size and activity levels are more strongly correlated with HHO periodicity in these animals. We argue that lemurs evolved this pattern due to selection for risk-averse life histories driven by the unpredictability of the environment in Madagascar. These results reinforce the idea that HHO influences life history evolution differently in response to specific ecological selection regimes. [ABSTRACT FROM AUTHOR]

Major achievements of primate evolution are skilled hand-object interaction and tool use, both in part dependent on parietal cortex expansion. We recorded spiking activity from macaque inferior parietal cortex during directional manipulation of an isometric tool, which required the application of hand forces to control a cursor's motion on a screen. In areas PFG/PF, the activity of 70%neurons was modulated by the hand force necessary to implement the desired target motion, reflecting an inverse model, rather than by the intended motion of the visual cursor (forward model). The population vector matched the direction and amplitude of the instantaneous force increments over time. When exposed to a new force condition, that obliged the monkey to change the force output to successfully bring the cursor to the final target, the activity of a consistent subpopulation of neurons changed in an orderly fashion and, at the end of a "Wash-out" session, retained memory of the new learned association, at the service of predictive control of force. Our findings suggest that areas PFG/PF represent a crucial node of the distributed control of hand force, by encoding instantaneous force variations and serving as a memory reservoir of hand dynamics required for object manipulation and tool use. This is coherent with previous studies in humans showing the following: (1) impaired adaptation to a new force field under TMS parietal perturbation; (2) defective control of direction of hand force after parietal lesion; and (3) fMRI activation of parietal cortex during object manipulation requiring control of fine hand forces. [ABSTRACT FROM AUTHOR]

Facultative bipedalism during load transport in nonhuman primates has been argued to be an important behavior potentially leading to the evolution of obligate, extended limb bipedalism. Understanding the biomechanics of such behavior may lead to insights about associated morphology, which may translate to interpretation of features in the fossil record. Some populations of bearded capuchin monkeys ( Sapajus libidinosus ) spontaneously carry heavy loads bipedally during foraging activities. This study provides the first data on all three components of ground reaction force for spontaneous bipedalism during load carriage in a nonhuman primate. Five individual S. libidinosus (mean body mass = 2.4 kg ± 0.96) were videorecorded during bipedalism while carrying a stone (0.93 kg) under natural conditions. A force plate was embedded in the path of the monkeys. Spatiotemporal and force data for all three components of the ground reaction force were recorded for 28 steps. Capuchins exhibited a mean vertical peak force per total weight (Vpk) for the hindlimb of 1.19 (sd = 0.13), consistent with those of unloaded capuchins in the laboratory and for other bipedal primates, including humans. Vertical force records suggest that capuchins, along with most nonhuman primates, maintain a relatively compliant leg during both unloaded and loaded locomotion. Like all other primates, loaded capuchins maintained laterally (outward) directed medio-lateral forces, presumably to stabilize side-to-side movements of the center of mass. Medio-lateral forces suggest that at near-running speeds dynamic stability diminishes the need to generate high lateral forces. Vertical force traces exhibited a measurable impact spike at foot contact in 85% of the steps recorded. An impact spike is common in human walking and running but has not been reported in other bipedal primates. This spike in humans is thought to lead to bone and cartilage damage. The earliest biped may have experienced similar impact spikes during bipedal locomotion, requiring compensatory behaviors or anatomical features. [ABSTRACT FROM AUTHOR]

Chewing on different food types is a demanding biological function. The classic assumption in studying the shape of feeding apparatuses is that animals are what they eat, meaning that adaptation to different food items accounts for most of their interspecific variation. Yet, a growing body of evidence points against this concept. We use the primate mandible as a model structure to investigate the complex interplay among shape, size, diet, and phylogeny. We find a weak but significant impact of diet on mandible shape variation in primates as a whole but not in anthropoids and catarrhines as tested in isolation. These clades mainly exhibit allometric shape changes, which are unrelated to diet. Diet is an important factor in the diversification of strepsirrhines and platyrrhines and a phylogenetic signal is detected in all primate clades. Peaks in morphological disparity occur during the Oligocene (between 37 and 25 Ma) supporting the notion that an adaptive radiation characterized the evolution of South American monkeys. In all primate clades, the evolution of mandible size is faster than its shape pointing to a strong effect of allometry on ecomorphological diversification in this group. [ABSTRACT FROM AUTHOR]

Background: The methyl-CpG Binding Protein two gene (MECP2) encodes a multifunctional protein comprising two isoforms involved in nuclear organization and regulation of splicing and mRNA template activity. This gene is normally expressed in all tissues, with a higher expression level in the brain during neuronal maturation. Loss of MECP2 function is the primary cause of Rett syndrome (RTT) in humans, a dominant, X-linked disorder dramatically affecting neural and motor development. Results: We investigated the molecular evolution of MECP2 in several primate taxa including 36 species in 16 genera of neotropical (platyrrhine) primates. The coding region of the MECP2_e2 isoform showed a high level of evolutionary conservation among humans and other primates, with amino acid substitutions in 14 codons and one in-frame insertion of a single serine codon, between codons 357 and 358, in Ateles paniscus. Most substitutions occurred in noncritical regions of MECP2 and the majority of the algorithms used for analyzing selection did not provide evidence of positive selection. Conversely, we found 48 sites under negative selection in different regions, 23 of which were consistently found by three different algorithms. Similar to an inverted Alu insert found previously in a lesser ape at a parallel location, one Alu insertion of approximately 300 bp in Cebus and Sapajus was found in intron 3. Phylogenetic reconstruction of the intron 3 data provided a topology that was coincident with the consensus arrangement of the primate taxa. RNAseq data in the neotropical primate Callimico goeldii revealed a novel transcript consisting of a noncontinuous region of the human-homologous intron 2 in this species; this transcript accounted for two putative polypeptides. Conclusions: Despite the remarkable evolutionary conservation of MECP2, one in-frame codon insertion was observed in A. paniscus, and one region of intron 3 was affected by a trans-specific Alu retrotransposition in two neotropical primate genera. Moreover, identification of novel MECP2 transcripts in Callimico suggests that part of a homologous human intronic region might be expressed, and that the potential open reading frame in this region might be a subject of interest in RTT patients who carry an apparently normal MECP2 sequence. [ABSTRACT FROM AUTHOR]

Previous analyses of dental size in Gigantopithecus blacki indicated marked sexual dimorphism and a trend towards increasing size through time. These studies were based on a sample of over 700 teeth from five localities excavated prior to 1990. Since then, 12 additional cave sites have been discovered in southern China, yielding hundreds of isolated teeth of G . blacki . Most of these sites are well dated by a combination of biochronology and absolute dating methods, so we now have a much better understanding of the chronology of G . blacki . Here, we reexamine the degree of sexual dimorphism and the question of dental size increase through time in G . blacki based on the expanded collections now available. Our results show that sexual dimorphism is not as marked as indicated in previous studies and confirm earlier analyses suggesting that the postcanine teeth of G. blacki tend to become larger through time from the beginning of the Early Pleistocene to the Middle Pleistocene. [ABSTRACT FROM AUTHOR]

This study tests the hypothesis that the third molars of Callimico goeldii represent a reversal in evolutionary tooth loss within the Callitrichinae. Loss of third molars is part of a suite of unusual characters that has been used to unite marmosets and tamarins in a clade to the exclusion of Callimico . However, molecular phylogenetic studies provide consistent support for the hypothesis that marmosets are more closely related to Callimico than to tamarins, raising the possibility that some or all of the features shared by marmosets and tamarins are homoplastic. Here, I use the binary-state speciation and extinction (BiSSE) model and a sample of 249 extant primate species to demonstrate that, given the shape of the primate phylogenetic tree and the distribution of character states in extant taxa, models in which M3 loss is constrained to be irreversible are much less likely than models in which reversals are allowed to occur. This result provides support for the idea that the last common ancestor of Callimico and marmosets was characterized by the two-molared phenotype. The M3s of Callimico therefore appear to be secondarily derived rather than plesiomorphic. This conclusion may also apply to the other apparently plesiomorphic traits found in Callimico . Hypotheses regarding the re-evolution of M3 in the callitrichine clade and the origin and maintenance of the two-molared phenotype are discussed. [ABSTRACT FROM AUTHOR]

The species Pseudoloris parvulus , identified in several Middle and Late Eocene European sites, was previously known in the Iberian Peninsula by a single mandible preserving P 4 –M 3 from Sossís (Southern Pyrenean Basins, northeastern Spain), described in the 1960s. Further field work at this Late Eocene site has led to the recovery of a large number of mammal remains, including the additional material of P. parvulus described in this paper. Some specimens of P. parvulus from this locality have also been recently found in the collections of the Naturhistorisches Museum Basel, Switzerland. The whole sample consists of 11 mandible fragments including several teeth, three upper dental series and nearly 80 isolated teeth including all of the dental elements, and represents the most complete sample of the genus described from the Iberian Peninsula. This abundant material allows us to provide an emended diagnosis for the species and to observe several directional changes in the dental morphology of the lineage including the species Pseudoloris saalae , Pseudoloris isabenae , Pseudoloris pyrenaicus and P. parvulus . These directional changes include the progressive reduction of the paraconid in the lower molars and the increase in size of the hypocone, metaconule and paraconule in the upper molars. Moreover, despite the overall resemblance among all of the samples ascribed to P. parvulus , we also recognize some differences, particularly an increase in size and better development of the hypocone from the oldest populations of the species, such as Le Bretou, to the most recent ones, like Sossís and Perrière. Therefore, this study sheds new light on the evolution of this genus, which inhabited Europe from the Middle Eocene to the Early Oligocene. [ABSTRACT FROM AUTHOR]

Despite significant progress in the structural and functional characterization of the human genome, understanding of the mechanisms underlying the genetic basis of human phenotypic uniqueness remains limited. Here, I report that transposable element-derived sequences, most notably LTR7/HERV-H, LTR5_Hs, and L1HS, harbor 99.8% of the candidate human-specific regulatory loci (HSRL) with putative transcription factor-binding sites in the genome of human embryonic stem cells (hESC). A total of 4,094 candidate HSRL display selective and site-specific binding of critical regulators (NANOG [Nanog homeobox], POU5F1 [POU class 5 homeobox 1], CCCTC-binding factor [CTCF], Lamin B1), and are preferentially located within the matrix of transcriptionally active DNA segments that are hypermethylated in hESC. hESC-specific NANOG-binding sites are enriched near the protein-coding genes regulating brain size, pluripotency long noncoding RNAs, hESC enhancers, and 5-hydroxymethylcytosine-harboring regions immediately adjacent to binding sites. Sequences of only 4.3% of hESC-specific NANOG-binding sites are present in Neanderthals' genome, suggesting that a majority of these regulatory elements emerged in Modern Humans. Comparisons of estimated creation rates of novel TF-binding sites revealed that there was 49.7-fold acceleration of creation rates of NANOG-binding sites in genomes of Chimpanzees compared with the mouse genomes and further 5.7-fold acceleration in genomes of Modern Humans compared with the Chimpanzees genomes. Preliminary estimates suggest that emergence of one novel NANOG-binding site detectable in hESC required 466 years of evolution. Pathway analysis of coding genes that have hESC-specific NANOG-binding sites within gene bodies or near gene boundaries revealed their association with physiological development and functions of nervous and cardiovascular systems, embryonic development, behavior, as well as development of a diverse spectrum of pathological conditions such as cancer, diseases of cardiovascular and reproductive systems, metabolic diseases, multiple neurological and psychological disorders. A proximity placement model is proposed explaining how a 33-47% excess of NANOG, CTCF, and POU5F1 proteins immobilized on a DNA scaffold may play a functional role at distal regulatory elements. [ABSTRACT FROM AUTHOR]

A non-adherence to a strict diurnal or nocturnal activity cycle is prevalent among mammals, including taxa of Lemuridae, but rare among other primates. While non-mutually exclusive ecological hypotheses attempted to explain the evolution of this activity, termed cathemerality, as either an old or a recent phenomenon, the scarcity of systematic data collected over 24 h limits our potential to explore its proximate and ultimate determinants. Among strepsirrhines, systematic studies involving only two lemurid genera ( Eulemur and Lemur) have recorded this activity pattern, while fewer quantitative observations are available for other taxa. If cathemerality could be shown in most members of Lemuridae despite their different ecological adaptations, this would support the hypothesis that this trait is basal and appeared early during lemurid evolution. Here, we investigated whether the folivorous southern bamboo lemur ( Hapalemur meridionalis) exhibits cathemeral activity, and determined which environmental factors influence its pattern. We deployed ten archival tags across four social groups to continuously record activity data over a 15-month period. This allowed us to generate a diurnal/nocturnal (DN) ratio for each 24-h period and assess their diel activity. Our data suggest that southern bamboo lemurs are cathemeral within Mandena; while climatic factors showed no influence, nocturnal activity increased with greater nocturnal luminance. Despite contrasting dietary niches, visual morphologies, and body sizes between Hapalemur, Eulemur, and Lemur, all three exhibit cathemerality and lunarphilia. The close phylogenetic proximity of these lemurids supports this flexible activity pattern as an ancestral trait that likely dates to the origin of the Lemuridae radiation. [ABSTRACT FROM AUTHOR]

ABSTRACT The notion of scala naturae dates back to thinkers such as Aristotle, who placed plants below animals and ranked the latter along a graded scale of complexity from 'lower' to 'higher' animals, such as humans. In the last decades, evolutionary biologists have tended to move from one extreme (i.e. the idea of scala naturae or the existence of a general evolutionary trend in complexity from 'lower' to 'higher' taxa, with Homo sapiens as the end stage) to the other, opposite, extreme (i.e. to avoid using terms such as 'phylogenetically basal' and 'anatomically plesiomorphic' taxa, which are seen as the undesired vestige of old teleological theories). The latter view tries to avoid any possible connotations with the original anthropocentric idea of a scala naturae crowned by man and, in that sense, it can be regarded as a more politically correct view. In the past years and months there has been renewed interest in these topics, which have been discussed in various papers and monographs that tend to subscribe, in general, to the points defended in the more politically correct view. Importantly, most evolutionary and phylogenetic studies of tetrapods and other vertebrates, and therefore most discussions on the scala naturae and related issues have been based on hard tissue and, more recently, on molecular data. Here we provide the first discussion of these topics based on a comparative myological study of all the major vertebrate clades and of myological cladistic and Bayesian phylogenetic analyses of bony fish and tetrapods, including Primates. We specifically ( i) contradict the notions of a scala naturae or evolutionary progressive trends leading to more complexity in 'higher' animals and culminating in Homo sapiens, and ( ii) stress that the refutation of these old notions does not necessarily mean that one should not keep using the terms 'phylogenetically basal' and particularly 'anatomically plesiomorphic' to refer to groups such as the urodeles within the Tetrapoda, or the strepsirrhines and lemurs within the Primates, for instance. This review will contribute to improving our understanding of these broad evolutionary issues and of the evolution of the vertebrate Bauplans, and hopefully will stimulate future phylogenetic, evolutionary and developmental studies of these clades. [ABSTRACT FROM AUTHOR]

Genetic diversity is a key parameter in population genetics and is important for understanding the process of evolution and for the development of appropriate conservation strategies. Recent advances in sequencing technology have enabled the measurement of genetic diversity of various organisms at the nucleotide level and on a genome-wide scale, yielding more precise estimates than were previously achievable. In this review, I have compiled and summarized the estimates of genetic diversity in humans and non-human primates based on recent genomewide studies. Although studies on population genetics demonstrated fluctuations in population sizes over time, general patterns have emerged. As shown previously, genetic diversity in humans is one of the lowest among primates; however, certain other primate species exhibit genetic diversity that is comparable to or even lower than that in humans. There exists greater than 10-fold variation in genetic diversity among primate species, and I found weak correlation with species fecundity but not with body or propagule size. I further discuss the potential evolutionary consequences of population size decline on the evolution of primate species. The level of genetic diversity negatively correlates with the ratio of nonsynonymous to synonymous polymorphisms in a population, suggesting that proportionally greater numbers of slightly deleterious mutations segregate in small rather than large populations. Although population size decline is likely to promote the fixation of slightly deleterious mutations, there are molecular mechanisms, such as compensatory mutations at various molecular levels, which may prevent fitness decline at the population level. The effects of slightly deleterious mutations from theoretical and empirical studies and their relevance to conservation biology are also discussed in this review. [ABSTRACT FROM AUTHOR]

The adaptive immune system uses V genes for antigen recognition. However, the evolutionary diversification and selection processes within and across species and orders remain poorly understood. Here, we studied the amino acid (AA) sequences obtained from the translated in-frame V exons of immunoglobulins (IG) and T cell receptors (TR) from 16 primate species whose genomes have been sequenced. Multi-species comparative analysis supports the hypothesis that V genes in the IG loci undergo birth/death processes, thereby permitting rapid adaptability over evolutionary time. We also show that multiple cladistic groupings exist in the TRA (35 clades) and TRB (25 clades) V gene loci and that each primate species typically contributes at least one V gene to each of these clades. The results demonstrate that IG V genes and TR V genes have quite different evolutionary pathways; multiple duplications can explain the IG loci results, while coevolutionary pressures can explain the phylogenetic results of the TR V gene loci. Our results suggest that there exist evolutionary relationships between V gene clades in the TRA and TRB loci. Due to the long-standing preservation of these clades, such genes may have specific and necessary roles for the viability of a species. [ABSTRACT FROM AUTHOR]

The Mx dynamin-like GTPases are key antiviral effector proteins of the type I and type III interferon (IFN) systems. They inhibit several different viruses by blocking early steps of the viral replication cycle. We focus on new structural and functional insights and discuss recent data revealing that human MxA (MX1) provides a safeguard against introduction of avian influenza A viruses (FLUAV) into the human population. The related human MxB (MX2) serves as restriction factor for HIV-1 and other primate lentiviruses. [ABSTRACT FROM AUTHOR]

Studies of primate cognition have conclusively shown that humans and apes share a range of basic cognitive abilities. As a corollary, these same studies have also focussed attention on what makes humans unique, and on when and how specifically human cognitive skills evolved. There is widespread agreement that a major distinguishing feature of the human mind is its capacity for causal reasoning. This paper argues that causal cognition originated with the use made of indirect natural signs by early hominins forced to adapt to variable late Miocene and early Pliocene environments; that early hominins evolved an innate tendency to search for such signs and infer their causes; that causal inference required the existence of incipient working memory; and that causal relationships were stored through being integrated into spatial maps to create increasingly complex causal models of the world. [ABSTRACT FROM AUTHOR]

ABSTRACT Teilhardina belgica is one of the earliest fossil primates ever recovered and the oldest fossil primate from Europe. As such, this taxon has often been hypothesized as a basal tarsiiform on the basis of its primitive dental formula with four premolars and a simplified molar cusp pattern. Until recently [see Rose et al.: Am J Phys Anthropol 146 (2011) 281-305; Gebo et al.: J Hum Evol 63 (2012) 205-218], little was known concerning its postcranial anatomy with the exception of its well-known tarsals. In this article, we describe additional postcranial elements for T. belgica and compare these with other tarsiiforms and with primitive adapiforms. The forelimb of T. belgica indicates an arboreal primate with prominent forearm musculature, good elbow rotational mobility, and a horizontal, rather than a vertical body posture. The lateral hand positions imply grasps adaptive for relatively large diameter supports given its small body size. The hand is long with very long fingers, especially the middle phalanges. The hindlimb indicates foot inversion capabilities, frequent leaping, arboreal quadrupedalism, climbing, and grasping. The long and well-muscled hallux can be coupled with long lateral phalanges to reconstruct a foot with long grasping digits. Our phyletic analysis indicates that we can identify several postcranial characteristics shared in common for stem primates as well as note several derived postcranial characters for Tarsiiformes. Am J Phys Anthropol 156:388-406, 2015. © 2014 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

Neuropeptide Y (NPY) plays a role in a variety of basic physiological functions and has also been implicated in regulating cognition, including learning and memory. A decrease in neocortical NPY has been reported for Alzheimer’s disease, schizophrenia, bipolar disorder, and depression, potentially contributing to associated cognitive deficits. The goal of the present analysis was to examine variation in neocortical NPY-immunoreactive axon and varicosity density among haplorhine primates (monkeys, apes, and humans). Stereologic methods were used to measure the ratios of NPY-expressing axon length density to total neuron density (ALv/Nv) and NPY-immunoreactive varicosity density to neuron density (Vv/Nv), as well as the mean varicosity spacing in neocortical areas 10, 24, 44, and 22 (Tpt) of humans, African great apes, New World monkeys, and Old World monkeys. Humans and great apes showed increased cortical NPY innervation relative to monkey species for ALv/Nv and Vv/Nv. Furthermore, humans and great apes displayed a conserved pattern of varicosity spacing across cortical areas and layers, with no differences between cortical layers or among cortical areas. These phylogenetic differences may be related to shared life history variables and may reflect specific cognitive abilities.

A correction is presented to the article "The first major primate extinction: An evaluation of paleoecological dynamics of North American stem primates using a homology free measure of tooth shape" by Kristen A. Prufrock, Doug M. Boyer and Mary T. Silcox which appeared in the previous issue.

Purpose: This article presents a novel concept of the evolution and, thus, the pathogenesis of uterine adenomyosis as well as peritoneal and peripheral endometriosis. Presently, no unifying denomination of this nosological entity exists., Methods: An extensive search of the literature on primate evolution was performed. This included comparative functional morphology with special focus on the evolution of the birthing process that fundamentally differs between the haplorrhine primates and most of the other eutherian mammals. The data were correlated with the results of own research on the pathophysiology of human archimetrosis and with the extant presentation of the disease., Results: The term Archimetrosis is suggested as a denomination of the nosological entity. Archimetrosis occurs in human females and also in subhuman primates. There are common features in the reproductive process of haplorrhine primates such as spontaneous ovulation and corpus luteum formation, spontaneous decidualization and menstruation. These have fused Müllerian ducts resulting in a uterus simplex. Following a usually singleton pregnancy, the fetus is delivered in the skull position. Some of these features are shared by other mammals, but not in that simultaneous fashion. In haplorrhine primates, with the stratum vasculare, a new myometrial layer has evolved during the time of the Cretaceous-Terrestrial Revolution (KTR) that subserves expulsion of the conceptus and externalization of menstrual debris in non-conceptive cycles. Hypercontractility of this layer has evolved as an advantage with respect to the survival of the mother and the birth of a living child during delivery and may be experienced as primary dysmenorrhea during menstruation. It may result in tissue injury by the sheer power of the contractions and possibly by the associated uterine ischemia. Moreover, the lesions at extra-uterine sites appear to be maintained by biomechanical stress., Conclusions: Since the pathogenesis of archimetrosis is connected with the evolution of the stratum vasculare, tissue injury and repair (TIAR) turns out to be the most parsimonious explanation for the development of the disease based on clinical, experimental and evolutionary evidence. Furthermore, a careful analysis of the published clinical data suggests that, in the risk population with uterine hypercontractility, the disease develops with a yet to be defined latency phase after the onset of the biomechanical injury. This opens a new avenue of prevention of the disease in potentially affected women that we consider to be primarily highly fertile., (© 2022. The Author(s).)

Evolutionary expansion and complexification of the primate cerebral cortex are largely linked to the emergence of the outer subventricular zone (OSVZ), a uniquely structured germinal zone that generates the expanded primate supragranular layers. The primate OSVZ departs from rodent germinal zones in that it includes a higher diversity of precursor types, inter-related in bidirectional non-hierarchical lineages. In addition, primate-specific regulatory mechanisms are operating in primate cortical precursors via the occurrence of novel miRNAs. Here, we propose that the origin and evolutionary importance of the OSVZ is related to genetic changes in multiple regulatory loops and that cell-cycle regulation is a favored target for evolutionary adaptation of the cortex. [ABSTRACT FROM AUTHOR]

A major evolutionary change in the lineage ancestral to humans, chimpanzee and gorilla (HCG) has been the embedding of the embryo into maternal tissue. Thus, the first layer of cells (trophoblast) to differentiate after fertilization must adapt to invade the uterus. Such event would likely leave signatures of positive selection at genes with roles in embryo implantation. Here, 163 pregnancy implantation genes are tested for evidence of adaptive diversification in the ancestral lineage to HCG. Two immune system genes, HLA-E and KIR2DL4 showed evidence of positive selection. Some of the positive selected sites involve amino acid substitution with predicted damaging effects on protein function, thus highlighting the possibility of antagonistic pleiotropic effects. Selection at a gene coding for a receptor expressed in uterine cells (KIR) that interacts with trophoblast human leukocyte antigen (HLA) genes suggests a main role for immunological adaptations in embryo deep invasion of the maternal endometrium. [ABSTRACT FROM AUTHOR]