Comparing vibrissal morphology and infraorbital foramen area in pinnipeds.

Pinniped vibrissae are well-adapted to sensing in an aquatic environment, by being morphologically diverse and more sensitive than those of terrestrial species. However, it is both challenging and time-consuming to measure vibrissal sensitivity in many species. In terrestrial species, the infraorbital foramen (IOF) area is associated with vibrissal sensitivity and increases with vibrissal number. While pinnipeds are thought to have large IOF areas, this has not yet been systematically measured before. We investigated vibrissal morphology, IOF area, and skull size in 16 species of pinniped and 12 terrestrial Carnivora species. Pinnipeds had significantly larger skulls and IOF areas, longer vibrissae, and fewer vibrissae than the other Carnivora species. IOF area and vibrissal number were correlated in Pinnipeds, just as they are in terrestrial mammals. However, despite pinnipeds having significantly fewer vibrissae than other Carnivora species, their IOF area was not smaller, which might be due to pinnipeds having vibrissae that are innervated more. We propose that investigating normalized IOF area per vibrissa will offer an alternative way to approximate gross individual vibrissal sensitivity in pinnipeds and other mammalian species. Our data show that many species of pinniped, and some species of felids, are likely to have strongly innervated individual vibrissae, since they have high values of normalized IOF area per vibrissa. We suggest that species that hunt moving prey items in the dark will have more sensitive and specialized vibrissae, especially as they have to integrate between individual vibrissal signals to calculate the direction of moving prey during hunting.

Good Vibrations: The Evolution of Whisking in Small Mammals.

While most mammals have whiskers, some tactile specialists-mainly small, nocturnal, and arboreal species-can actively move their whiskers in a symmetrical, cyclic movement called whisking. Whisking enables mammals to rapidly, tactually scan their environment to efficiently guide locomotion and foraging in complex habitats. The muscle architecture that enables whisking is preserved from marsupials to primates, prompting researchers to suggest that a common ancestor might have had moveable whiskers. Studying the evolution of whisker touch sensing is difficult, and we suggest that measuring an aspect of skull morphology that correlates with whisking would enable comparisons between extinct and extant mammals. We find that whisking mammals have larger infraorbital foramen (IOF) areas, which indicates larger infraorbital nerves and an increase in sensory acuity. While this relationship is quite variable and IOF area cannot be used to solely predict the presence of whisking, whisking mammals all have large IOF areas. Generally, this pattern holds true regardless of an animal's substrate preferences or activity patterns. Data from fossil mammals and ancestral character state reconstruction and tracing techniques for extant mammals suggest that whisking is not the ancestral state for therian mammals. Instead, whisking appears to have evolved independently as many as seven times across the clades Marsupialia, Afrosoricida, Eulipotyphla, and Rodentia, with Xenarthra the only placental superordinal clade lacking whisking species. However, the term whisking only captures symmetrical and rhythmic movements of the whiskers, rather than all possible whisker movements, and early mammals may still have had moveable whiskers. Anat Rec, 2018. © 2018 American Association for Anatomy.

Scaling of Anatomically Derived Maximal Bite Force in Primates.

By combining muscle architectural data with biomechanical variables relating to the jaw, we produce anatomically derived maximum bite force estimations for 23 species of catarrhine and platyrrhine primates. We investigate how bite force scales across the sample as a whole (and within each parvorder) relative to two size proxies, body mass and cranial geometric mean, and the effect of diet upon bite force. Bite force is estimated at three representative bite points along the dental row: the first maxillary incisor, canine, and third-most mesial paracone. We modeled bite force by combining calculated physiological cross-sectional area of the jaw adductors from Hartstone-Rose et al. [Anat Rec 301 (2018) 311-324] with osteological measurements of lever- and load-arm lengths from the same specimens [Hartstone-Rose et al., Anat Rec 295 (2012) 1336-1351]. Bite force scales with positive allometry relative to cranial geometric mean across our entire sample and tends toward positive allometry relative to body mass. Bite force tends toward positive allometry within platyrrhines but scales isometrically within catarrhines. There was no statistically significant scaling difference with diet. Our findings imply an absence of a dietary signal in the scaling of bite force, a result that differs from the scaling of physiological cross-sectional area alone. That is, although previous studies have found a dietary signal in the muscle fiber architecture in these species, when these are combined with their leverages, that signal is undetectable. On the parvorder level, our data also demonstrate that the platyrrhine masticatory system appears more mechanically advantageous than that of catarrhines. Anat Rec, 2019. © 2019 American Association for Anatomy Anat Rec, 303:2026-2035, 2020. © 2019 American Association for Anatomy.

Vertical Clinging and Leaping Ahead: How Bamboo Has Shaped the Anatomy and Physiology of Hapalemur.

Hapalemur sps. and Prolemur simus (bamboo lemurs, collectively) stand out from the relatively homogeneous lemurids because they are bamboo feeders and vertical clingers and leapers. This unique diet presents equally unique challenges, like its verticality, toughness, and toxicity. The bamboo lemurs share the generalized anatomy of the other lemurids, but also display some well-documented skeletal adaptations, perhaps to overcome the problems presented by their specialization. Soft-tissue adaptations, however, remain largely unexplored. Explored here are possible soft-tissue adaptations in Hapalemur griseus. We compare H. griseus with other lemurids, Propithecus, Galago, Tarsier, and a tree shrew. Based on the available anatomical and physiological data, we hypothesize that Hapalemur and Prolemur species will have differences in hindlimb morphology when compared with other lemurids. We predict that H. griseus will have more hindlimb muscle mass and will amplify muscle mass differences with increased type II muscle fibers. Relative hindlimb muscle mass in H. griseus is less than other prosimians sampled, yet relative sural muscle mass is significantly heavier (P < 0.01) in H. griseus. Results show that the soleus muscle of H. griseus has a higher amount of type II (fast) fibers in plantarflexors. These findings indicate although H. griseus shares some generalized lemurid morphology, its diet of bamboo may have pushed this generalized lemurid to an anatomical extreme. We suspect additional bamboo-specific adaptations in their anatomy and physiology will be uncovered with further examination into the anatomy of the bamboo lemurs. Anat Rec, 2019. © 2019 Wiley Periodicals, Inc. Anat Rec, 303:295-307, 2020. © 2019 American Association for Anatomy.

The ligamentum teres femoris in orangutans.

OBJECTIVES: It is widely viewed that orangutans lack a ligamentum teres femoris (LTF) inserting on the femoral head because orangutans lack a distinct fovea capitis. Orangutans employ acrobatic quadrumanous clambering that requires a high level of hip joint mobility, and the absence of an LTF is believed to be an adaptation to increase hip mobility. However, there are conflicting reports in the literature about whether there may be a different LTF configuration in orangutans, perhaps with a ligament inserting on the femoral neck instead. Here we perform a dissection-based study of orangutan hip joints, assess the soft tissue and hard tissue correlates of the orangutan LTF, and histologically examination the LTF to evaluate whether it is homologous to that found in other hominoids. MATERIALS AND METHODS: The hip joints from six orangutans were dissected. In the two orangutans with an LTF passing to the femoral head, the LTF was assessed histologically. Skeletonized femora (n=56) in osteological repositories were examined for evidence of a foveal pit. RESULTS: We observed an LTF in two of the three infant orangutans but not in the sub-adult or adult specimens. Histological examination of the infant LTF shows a distinct artery coursing through the LTF to the head of the femur. One percent of orangutan femora present with a foveal scar, but no pit, on the femoral head. DISCUSSION: Despite being absent in adults, the LTF is present in at least some orangutans during infancy. We suggest that the LTF maintains a role in blood supply to the femoral head early in life. Because the LTF can limit hip mobility, this may explain why the LTF may be lost as an orangutan ages and gains locomotor independence. These findings enhance our understanding of orangutan hip morphology and underscore the need for future soft tissue investigations.

How the Brain May Have Shaped Muscle Anatomy and Physiology: A Preliminary Study.

Skeletal muscle fibers are often used to evaluate functional differences in locomotion. However, because there are energetic differences among muscle fiber cells, muscle fiber composition could be used to address evolutionary questions about energetics. Skeletal muscle is composed of two main types of fibers: Type I and II. The difference between the two can be reduced to how these muscle cells use oxygen and glucose. Type I fibers convert glucose to ATP using oxygen, while Type II fibers rely primarily on anaerobic metabolic processes. The expensive tissue hypothesis (ETH) proposes that the energetic demands imposed on the body by the brain result in a reduction in other expensive tissues (e.g., gastrointestinal tract). The original ETH dismisses the energetic demands of skeletal muscle, despite skeletal muscle being (1) an expensive tissue when active and (2) in direct competition for glucose with the brain. Based on these observations we hypothesize that larger brained primates will have relatively less muscle mass and a decrease in Type I fibers. As part of a larger study to test this hypothesis, we present data from 10 species of primates. We collected body mass, muscle mass, and biopsied four muscles from each specimen for histological procedures. We collected endocranial volumes from the literature. Using immunohistochemistry, a muscle fiber composition profile was created for each species sampled. Results show that larger brained primates have less muscle and fewer Type I fibers than primates with smaller brains. Results clarify the relationship between muscle mass and brain mass and illustrate how muscle mass could be used to address energetic questions. Anat Rec, 301:528-537, 2018. © 2018 Wiley Periodicals, Inc.

Relative tooth size at birth in primates: Life history correlates.

OBJECTIVES: Dental eruption schedules have been closely linked to life history variables. Here we examine a sample of 50 perinatal primates (28 species) to determine whether life history traits correlate with relative tooth size at birth. MATERIALS AND METHODS: Newborn primates were studied using serial histological sectioning. Volumes of deciduous premolars (dp2 -dp4 ), replacement teeth (if any), and permanent molars (M1-2/3 ) of the upper jaw were measured and residuals from cranial length were calculated with least squares regressions to obtain relative dental volumes (RDVs). RESULTS: Relative dental volumes of deciduous or permanent teeth have an unclear relationship with relative neonatal mass in all primates. Relative palatal length (RPL), used as a proxy for midfacial size, is significantly, positively correlated with larger deciduous and permanent postcanine teeth. However, when strepsirrhines alone are examined, larger RPL is correlated with smaller RDV of permanent teeth. In the full sample, RDVs of deciduous premolars are significantly negatively correlated with relative gestation length (RGL), but have no clear relationship with relative weaning age. RDVs of molars lack a clear relationship with RGL; later weaning is associated with larger molar RDV, although correlations are not significant. When strepsirrhines alone are analyzed, clearer trends are present: longer gestations or later weaning are associated with smaller deciduous and larger permanent postcanine teeth (only gestational length correlations are significant). DISCUSSION: Our results indicate a broad trend that primates with the shortest RGLs precociously develop deciduous teeth; in strepsirrhines, the opposite trend is seen for permanent molars. Anthropoids delay growth of permanent teeth, while strepsirrhines with short RGLs are growing replacement teeth concurrently. A comparison of neonatal volumes with existing information on extent of cusp mineralization indicates that growth of tooth germs and cusp mineralization may be selected for independently.

A comparative analysis of infraorbital foramen size in Paleogene euarchontans.

The size of the infraorbital foramen (IOF) is correlated with the size of the infraorbital nerve and number of mystacial vibrissae in mammals. Accordingly, IOF cross-sectional area has been used to infer both the rostral mechanoreceptive acuity and phylogenetic relationships of extinct crown primates and plesiadapiforms. Among living mammals, extant primates, scandentians, and dermopterans (Euarchonta) exhibit smaller IOF cross-sectional areas than most other mammals. Here we assess whether fossil adapoids, omomyoids, and plesiadapiforms show a reduction in relative IOF area similar to that characterizing extant euarchontans. The IOFs of 12 adapoid, 7 omomyoid, 15 plesiadapiform, and 3 fossil gliran species were measured and compared to a diverse extant mammalian sample. These data demonstrate that adapoids and omomyoids have IOFs that are similar in relative size to those of extant euarchontans. Conversely, IOFs of plesiadapiforms are on average about twice as large as those of extant euarchontans and are more comparable in size to those of extant non-euarchontan mammals. These results indicate that crown primates share a derived reduction in relative IOF size with treeshrews and colugos. Accordingly, a decreased reliance on the muzzle and an increased reliance on the hands for environmental exploration may have first evolved in the euarchontan stem lineage. However, the relatively large IOFs of plesiadapiforms imply a continued reliance on the muzzle for close exploration of objects. This finding may indicate that either parallel evolutionary decreases in IOF size occurred within Euarchonta or that plesiadapiforms lie outside the euarchontan crown group.

Dietary correlates associated with the mental foramen in primates: implications for interpreting the fossil record.

The mandibular nerve is a sensory and motor nerve that innervates the muscles of mastication, the lower dentition, and the lower lip and surrounding structures. Although this nerve contains both efferent and afferent fibers, the mental nerve, a terminal branch of the mandibular nerve, is a strictly sensory nerve that exits the mental foramen and innervates the lower lip, the skin overlaying the mandible, and the oral mucosa around the mandible. Osteological foramina are often used as proxies for nerve cross section area and they often correlate well with some aspect of a primate's ecology (e.g., optic foramen and visual acuity). The primary objective of this study is to explore the correlation between the mental foramen and dietary preference among primates. The mental foramen of 40 primate species (n = 180) was measured from 3-D surface models of the mandible. Both conventional and phylogenetic tests indicate that although frugivores have larger mental foramina than folivores, the differences were not significant. These results show that while structures like the infraorbital foramen correlate well with diet and touch sensitivity, the mental foramen does not. Based on these findings, the mental foramen is not a suggested morphological character for interpreting of the fossil record. J. Morphol. 277:978-985, 2016. © 2016 Wiley Periodicals, Inc.

Does the primate pattern hold up? Testing the functional significance of infraorbital foramen size variation among marsupials.

OBJECTIVES: The relative size of the infraorbital foramen (IOF) has been used to infer the ecology of extinct primates for several decades. Primates have relatively smaller IOFs than most other mammals, which may result from the fact that they pre-process and manipulate food with their hands rather than their muzzles. In primates, relative IOF area co-varies with diet, where insectivores and folivores have relatively smaller IOFs than frugivores. We wanted to determine whether the observed patterns associated with IOF variation hold across other orders. MATERIALS AND METHODS: We examined how relative IOF area differs among marsupials occupying different ecological niches. Marsupials were chosen because they converge with primates in both ecology and morphology, but unlike primates, some marsupials approach and pre-process foods only with their muzzles. We measured IOF area and cranial lengths from 72 marsupial species, and behavioral feeding data were obtained from a subset of this sample (N = 20). RESULTS: Relative IOF area did not vary significantly between substrate preferences. However, relative IOF area differed significantly by diet category (P < 0.001). Species that specialize in feeding on non-grassy leaves have significantly smaller relative IOF areas than species which primarily feed on grasses, insects, vertebrates, or some combination thereof. Behavioral analyses support that folivorous marsupials approach and remove food with the hands more often than marsupials from other dietary groups. DISCUSSION: Results suggest that relatively small IOF area may reflect increased reliance on the hands while feeding, and that relative IOF size can be used as an indicator of feeding behavior.

Dental maturation, eruption, and gingival emergence in the upper jaw of newborn primates.

In this report we provide data on dental eruption and tooth germ maturation at birth in a large sample constituting the broadest array of non-human primates studied to date. Over 100 perinatal primates, obtained from natural captive deaths, were screened for characteristics indicating premature birth, and were subsequently studied using a combination of histology and micro-CT. Results reveal one probable unifying characteristic of living primates: relatively advanced maturation of deciduous teeth and M1 at birth. Beyond this, there is great diversity in the status of tooth eruption and maturation (dental stage) in the newborn primate. Contrasting strategies in producing a masticatory battery are already apparent at birth in strepsirrhines and anthropoids. Results show that dental maturation and eruption schedules are potentially independently co-opted as different strategies for attaining feeding independence. The most common strategy in strepsirrhines is accelerating eruption and the maturation of the permanent dentition, including replacement teeth. Anthropoids, with only few exceptions, accelerate mineralization of the deciduous teeth, while delaying development of all permanent teeth except M1. These results also show that no living primate resembles the altricial tree shrew (Tupaia) in dental development. Our preliminary observations suggest that ecological explanations, such as diet, provide an explanation for certain morphological variations at birth. These results confirm previous work on perinatal indriids indicating that these and other primates telegraph their feeding adaptations well before masticatory anatomy is functional. Quantitative analyses are required to decipher specific dietary and other influences on dental size and maturation in the newborn primate.

The vomeronasal organ of Lemur catta.

The vomeronasal organ (VNO), also known as the Jacobson's organ, is a bilateral chemosensory organ found at the base of the nasal cavity specialized for the detection of higher-molecular weight (non-volatile) chemostimuli. It has been linked to pheromone detection. The VNO has been well studied in nocturnal lemurs and lorises, but poorly studied in diurnal/cathemeral species despite the large repertoire of olfactory behaviors noted in species such as Lemur catta. Here, the VNO and associated structures were studied microanatomically in one adult female and one adult male L. catta. Traditional and immunohistochemical procedures demonstrate the VNO epithelium consists of multiple rows of sensory neurons. Immunoreactivity to Growth-associated protein 43 (GAP43) indicates the VNO is postnatally neurogenic. In volume, the VNO neuroepithelium scales similarly to palatal length compared to nocturnal strepsirrhines. Numerous taste buds present at the oral opening to the nasopalatine duct, with which the VNO communicates, provide an additional (or alternative) explanation for the flehmen behavior that has been observed in this species. The VNO of L. catta is shown to be microanatomically comparable to that of nocturnal strepsirrhines. Like nocturnal strepsirrhines, the VNO of L. catta may be functional in the reception of high-molecular weight secretions.

Caudal vertebral body articular surface morphology correlates with functional tail use in anthropoid primates.

Prehensile tails, capable of suspending the entire body weight of an animal, have evolved in parallel in New World monkeys (Platyrrhini): once in the Atelinae (Alouatta, Ateles, Brachyteles, Lagothrix), and once in the Cebinae (Cebus, Sapajus). Structurally, the prehensile tails of atelines and cebines share morphological features that distinguish them from nonprehensile tails, including longer proximal tail regions, well-developed hemal processes, robust caudal vertebrae resistant to higher torsional and bending stresses, and caudal musculature capable of producing higher contractile forces. The functional significance of shape variation in the articular surfaces of caudal vertebral bodies, however, is relatively less well understood. Given that tail use differs considerably among prehensile and nonprehensile anthropoids, it is reasonable to predict that caudal vertebral body articular surface area and shape will respond to use-specific patterns of mechanical loading. We examine the potential for intervertebral articular surface contour curvature and relative surface area to discriminate between prehensile-tailed and nonprehensile-tailed platyrrhines and cercopithecoids. The proximal and distal intervertebral articular surfaces of the first (Ca1), transitional and longest caudal vertebrae were examined for individuals representing 10 anthropoid taxa with differential patterns of tail-use. Study results reveal significant morphological differences consistent with the functional demands of unique patterns of tail use for all vertebral elements sampled. Prehensile-tailed platyrrhines that more frequently use their tails in suspension (atelines) had significantly larger and more convex intervertebral articular surfaces than all nonprehensile-tailed anthropoids examined here, although the intervertebral articular surface contour curvatures of large, terrestrial cercopithecoids (i.e., Papio sp.) converge on the ateline condition. Prehensile-tailed platyrrhines that more often use their tails in tripodal bracing postures (cebines) are morphologically intermediate between atelines and nonprehensile tailed anthropoids.

The interpretive power of infraorbital foramen area in making dietary inferences in extant apes.

The infraorbital foramen (IOF) is located below the orbit and transmits the sensory infraorbital nerve (ION) to mechanoreceptors located throughout the maxillary region. The size of the IOF correlates with the size of the ION; thus, the IOF appears to indicate relative touch sensitivity of maxillary region. In primates, IOF size correlates well with diet. Frugivores have relatively larger IOFs than folivores or insectivores because fruit handling/processing requires increased touch sensitivity. However, it is unknown if the IOF can be used to detect subtle dietary differences among closely related hominoid species. Hominoids are traditionally grouped into broad dietary categories, despite the fact that hominoid diets are remarkably diverse. This study examines whether relative IOF size is capable of differentiating among the dietary preferences of closely related species with overlapping, yet divergent diets. We measured IOF area in Hylobates lar, Symphalangus syndactulus, Pongo pygmaeus spp., Pan troglodytes, Gorilla gorilla, Gorilla beringei graueri, and Gorilla beringei beringei. We classified each species as a dedicated folivore, mixed folivore/frugivore, soft object frugivore, or hard object frugivore. The IOF is documented to be larger in more frugivorous species and smaller in more folivorous taxa. Interestingly, G.b. beringei, had the largest relative IOF of any gorilla, despite being a dedicated folivore. G.b. beringei does have unique food processing behavior that relies heavily on maxillary mechanoreception, thus this finding is not entirely unsupported behaviorally. The results of this study provide evidence that the IOF is an informative feature in interpretations of fossil apes.

Comparative histomorphology of intrinsic vibrissa musculature among primates: implications for the evolution of sensory ecology and "face touch".

Macrovibrissae are specialized tactile sensory hairs present in most mammalian orders, used in maxillary mechanoreception or "face touch." Some mammals have highly organized vibrissae and are able to "whisk" them. Movement of vibrissae is influenced by intrinsic vibrissa musculature, striated muscle bands that attach directly to the vibrissa capsule. It is unclear if primates have organized vibrissae or intrinsic vibrissa musculature and it is uncertain if they can move their vibrissae. The present study used histomorphological techniques to compare vibrissae among 19 primates and seven non-primate mammalian taxa. Upper lips of these mammals were sectioned and processed for histochemical analysis. While controlling for phylogenetic effects the following hypotheses were tested: 1) mammals with well-organized vibrissae possess intrinsic vibrissa musculature and 2) intrinsic vibrissa musculature is best developed in nocturnal, arboreal taxa. Our qualitative analyses show that only arboreal, nocturnal prosimians possess intrinsic musculature. Not all taxa that possessed organized vibrissae had intrinsic vibrissa musculature. Phylogenetic comparative analyses revealed a 70% probability that stem mammals, primates, and haplorhines possessed intrinsic vibrissa musculature and well-organized vibrissae. These two traits most likely coevolved according to a discrete phylogenetic analysis. These results indicate that nocturnal, arboreal primates have the potential to more actively use their vibrissae in spatial recognition and navigation tasks than diurnal, more terrestrial species, but there is a clear phylogenetic signal involved in the evolution of primate vibrissae and "face touch."

Eye size at birth in prosimian primates: life history correlates and growth patterns.

BACKGROUND: Primates have large eyes relative to head size, which profoundly influence the ontogenetic emergence of facial form. However, growth of the primate eye is only understood in a narrow taxonomic perspective, with information biased toward anthropoids. METHODOLOGY/PRINCIPAL FINDINGS: We measured eye and bony orbit size in perinatal prosimian primates (17 strepsirrhine taxa and Tarsius syrichta) to infer the extent of prenatal as compared to postnatal eye growth. In addition, multiple linear regression was used to detect relationships of relative eye and orbit diameter to life history variables. ANOVA was used to determine if eye size differed according to activity pattern. In most of the species, eye diameter at birth measures more than half of that for adults. Two exceptions include Nycticebus and Tarsius, in which more than half of eye diameter growth occurs postnatally. Ratios of neonate/adult eye and orbit diameters indicate prenatal growth of the eye is actually more rapid than that of the orbit. For example, mean neonatal transverse eye diameter is 57.5% of the adult value (excluding Nycticebus and Tarsius), compared to 50.8% for orbital diameter. If Nycticebus is excluded, relative gestation age has a significant positive correlation with relative eye diameter in strepsirrhines, explaining 59% of the variance in relative transverse eye diameter. No significant differences were found among species with different activity patterns. CONCLUSIONS/SIGNIFICANCE: The primate developmental strategy of relatively long gestations is probably tied to an extended period of neural development, and this principle appears to apply to eye growth as well. Our findings indicate that growth rates of the eye and bony orbit are disassociated, with eyes growing faster prenatally, and the growth rate of the bony orbit exceeding that of the eyes after birth. Some well-documented patterns of orbital morphology in adult primates, such as the enlarged orbits of nocturnal species, mainly emerge during postnatal development.

Muscle mass scaling in primates: an energetic and ecological perspective.

Body composition is known to vary dramatically among mammals, even in closely related species, yet this issue has never been systematically investigated. Here, we examine differences in muscle mass scaling among mammals, and explore how primate body composition compares to that of nonprimate mammals. We use a literature-based sample of eutherian and metatherian mammals, and combine this with new dissection-based data on muscularity in a variety of strepsirrhine primates and the haplorhine, Tarsius syrichta. Our results indicate an isometric scaling relationship between total muscle mass and total body mass across mammals. However, we documented substantial variation in muscularity in mammals (21-61% of total body mass), which can be seen both within and between taxonomic groups. We also found that primates are under-muscled when compared to other mammals. This difference in body composition may in part reflect the functional consequences of arboreality, as arboreal species have significantly lower levels of muscularity than terrestrial species.

Mechanoreceptivity of prehensile tail skin varies between ateline and cebine primates.

Prehensile tails evolved independently twice in primates: once in the ateline subfamily of platyrrhine primates and once in the genus Cebus. Structurally, the prehensile tails of atelines and Cebus share morphological features distinguishing them from nonprehensile tails (e.g., robust and strong caudal vertebrae, well developed lateral tail musculature, etc.). However, because of their independent evolutionary histories, the prehensile tails of atelines exhibit some differences from the Cebus prehensile tail. Ateline tails are relatively longer than those of Cebus, and they have less well-developed extensor compartment musculature. However, perhaps the most obvious difference is the distinctive hairless friction pad on the ventrodistal surface of the ateline tail; the tail of Cebus is completely covered in hair. This study documents the presence of four epicritic histologic mechanoreceptors in the friction pad of atelines: Meissner's corpuscles, Pacinian corpuscles, Ruffini corpuscles, and Merkel discs. Ruffini corpuscles and Merkel cells were also identified in the ventrodistal skin of the Cebus tail. However, Meissner's and Pacinian corpuscles (not typically associated with hairy skin) were not found in Cebus. Cebus was also compared to its closest living sister taxon, nonprehensile-tailed Saimiri, in which genus only Ruffini corpuscles are observed (no Merkel discs). The differences in mechanoreceptor type and morphology are attributed to the contrasting behavioral and tactile demands of the tail as it is used in posture and locomotion, which also distinguishes atelines from Cebus.

Behavioral and ecological consequences of sex-based differences in gustatory anatomy in Cebus apella.

Fungiform papillae (FPs) are the only gustatory structures on the anterior tongue. Taste buds (TBs), which are located in FPs, house taste receptors. Each TB has a taste pore (TP) by which tastants are transmitted. In humans, FP and TB densities correlate with taste sensitivity and food preferences. Females have higher FP densities than males in Homo, Pan, and Cebus. Homo, Pan, and Cebus also have larger brains, slower ontogenetic development, and higher maternal investment in offspring compared to most primates. An increase in maternal investment places intense pressure on females to 1) obtain high-quality foods, and 2) detect potential toxins at low levels. This study examines sex differences in FPs and TPs (a TB surrogate) in 11 Cebus apella to test the hypothesis that higher FP density in females may be an adaptation specific to reproductive strategies of females. Tongues were imaged using an environmental scanning electron microscope; from these images FP surface area, FP density, TP count, and TP densities were calculated. We found that there were no significant differences between males and females in the number of TPs per FP. However, we did find that females do have larger FP surface areas and higher FP densities than males. The anatomical evidence indicates that females may have greater taste sensitivity than males because females have more FP than males. Future research on food preference and selection in Cebus is expected to show sex-specific behaviors similar to those observed in Homo and Pan.

Anatomical Correlates to Nectar Feeding among the Strepsirrhines of Madagascar: Implications for Interpreting the Fossil Record.

One possible ecological scenario for the origin of primates is the archaic pollination and coevolution hypothesis. Its proponents contend that the consumption of nectar by some early primates and the resulting cross-pollination is an example of coevolution that drove adaptive radiations in some primates. This hypothesis is perhaps ecologically sound, but it lacks the morphology-behavior links that would allow us to test it using the fossil record. Here we attempt to identify cranial adaptations to nectar feeding among the strepsirrhines of Madagascar in order to provide such links. Many Malagasy strepsirrhines are considered effective cross-pollinators of the flowers they feed from, and nectar consumption represents as much as 75% of total feeding time. Previous studies identified skeletal correlates to nectar feeding in the crania of nonprimate mammals; from these, nine cranial measurements were chosen to be the focus of the present study. Results indicate that Cheirogaleus, Varecia, and Eulemur mirror other nectar-feeding mammals in having elongated crania and/or muzzles. These strepsirrhines might be effective cross-pollinators, lending support to the coevolution hypothesis.