W. Scott McGraw

Hard-object feeding in sooty mangabeys (Cercocebus atys) and interpretation of early hominin feeding ecology.

Morphology of the dentofacial complex of early hominins has figured prominently in the inference of their dietary adaptations. Recent theoretical analysis of craniofacial morphology of Australopithecus africanus proposes that skull form in this taxon represents adaptation to feeding on large, hard objects. A modern analog for this specific dietary specialization is provided by the West African sooty mangabey, Cercocebus atys. This species habitually feeds on the large, exceptionally hard nuts of Sacoglottis gabonensis, stereotypically crushing the seed casings using their premolars and molars. This type of behavior has been inferred for A. africanus based on mathematical stress analysis and aspects of dental wear and morphology. While postcanine megadontia, premolar enlargement and thick molar enamel characterize both A. africanus and C. atys, these features are not universally associated with durophagy among living anthropoids. Occlusal microwear analysis reveals complex microwear textures in C. atys unlike those observed in A. africanus, but more closely resembling textures observed in Paranthropus robustus. Since sooty mangabeys process hard objects in a manner similar to that proposed for A. africanus, yet do so without the craniofacial buttressing characteristic of this hominin, it follows that derived features of the australopith skull are sufficient but not necessary for the consumption of large, hard objects. The adaptive significance of australopith craniofacial morphology may instead be related to the toughness, rather than the hardness, of ingested foods.

Feeding, diet, and jaw form in West African Colobus and Procolobus

The functional link between mandibular morphology and masticatory stress has been documented by both experimental and comparative investigation. Somewhat more tenuous is the purported connection between dietary variation and the form of the jaws in primates. Several factors complicate the inference of such a connection, including anecdotal or incomplete dietary data from field studies and allometric effects on skeletal form that may have little to do with diet per se. We compared the jaws of sympatric colobines from West Africa to test the effect of diet on mandibular form. Procolobus badius and Colobus polykomos occupy the same habitat yet differ in diet primarily due to the exploitation of hard seeds by C. polykomos. The fact that the two taxa are comparable in body size also obviates the need for allometric qualifications. Colobus polykomos is expected to possess more robust mandibular corpora than Procolobus badius. In fact, the jaws of Colobus polykomos do not differ consistently from those of Procolobus badius in terms of biomechanical function. This apparent failure of mandibular morphology to reflect differences in diet and feeding behavior may be due to a variety of factors. We suspect that functional demands related to canine tooth support are contributing to obliteration of the expected biomechanical signal. Successful prediction of dietary effects on mandibular form requires consideration of competing structural and functional demands. The influence of diet on mandibular corporeal morphology is not equivalent across different primate species.

Sex and age differences in the diet and ingestive behaviors of sooty mangabeys (Cercocebus atys) in the Tai forest, Ivory coast

Members of the Cercocebus-Mandrillus clade are united by several morphological features, including expanded premolars which are argued to be associated with a preponderance of hard objects in the diet. We test the association between premolar expansion and hard object feeding by examining how different dental regions are used during food processing. We examined the diet and oral processing activities of sooty mangabeys (Cercocebus atys) in the Ivory Coasts Tai forest from August 2008 to September 2009. In addition to compiling diet profiles, we recorded the frequency that individuals performed four activities: 1) incising, 2) canine puncturing/scraping, 3) postcanine crushing (i.e., isometric biting), and 4) routine mastication (chewing cycles). Sooty mangabeys have a relatively narrow diet that consists largely of nuts/seeds, fruits, and invertebrates. While there are age and sex differences in diet, the most frequently consumed foods are similar across age and sex classes. The most frequently consumed foods are seeds of Sacoglottis gabonensis which are the hardest items in the sooty mangabey diet. Patterns of ingestive behavior vary with food type, but adults and nonadults (excluding infants dependent on mothers) of both sexes process similar foods. Premolar expansion in Cercocebus atys is associated with powerful crushing of hard objects of specific size and durophagy is a constant feature of sooty mangabey feeding ecology throughout ontogeny.

Association of Terrestrial Mangabeys (Cercocebus atys) with Arboreal Monkeys: Experimental Evidence for the Effects of Reduced Ground Predator Pressure on Habitat Use

We examined the behavior of two arboreal monkeys—Piliocolobus badius (western red colobus) and Cercopithecus diana (Diana monkey)—in the presence and absence of a third, predominately terrestrial monkey, Cercocebus atys (sooty mangabey) in the Ivory Coasts Taï Forest. Via experimental data, we show that sooty mangabeys are effective sentinels for ground predators. Then we compared strata use of red colobus and Diana monkeys in the presence and absence of mangabeys. Our data indicate that red colobus and Diana monkeys descend to low forest levels and to the forest floor significantly more often when mangabeys are spatially intermingled with them, i.e. in polyspecific associations. The niche extension may provide some foraging advantage, especially for red colobus. We identified the specific causal agent—relaxed ground predator pressure—of a temporary shift in habitat use. In this instance, the presence of one species (Cercocebus atys) leads to the temporary expansion of a niche normally avoided by others Piliocolobus badius and Cercopithecus diana.

The Monkeys of the Taï forest : an African primate community

1: The monkeys of the Tai Forest: an introduction W. S. McGraw and K. Zuberbuhler, Part I. Social Behavior: 2. The social system of guenons P. Buzzard and W .Eckardt 3: How small-scale differences in food competition lead to different social systems in three closely related sympatric colobines A. H. Korstjens, K. Bergman, C.Deffernez, M. Krebs, E. C. Nijssen, BAM van Oirschot, C Paukert, E. P. Schippers 4. The structure of social relationships among sooty mangabeys in Tai F. Range, T. Forderer, Y. Meystre, C. Benetton, and C. Fruteau Part II. Anti-Predation Strategies: 5. Interactions between leopard and monkeys K. Zuberbuhler and D. Jenny 6. Interactions between red colobus and chimpanzees R. Bshary 7. Interactions between African crowned eagles and their primate prey community S. Shultz and S. Thomsett 8. Semantic information in alarm calls K. Zuberbuhler Part III. Habitat Use: 9. Positional behavior and habitat use of Tai Forest monkeys W. S. McGraw Part IV. Conservation: 10. Can monkey behavior be used as an indicator for poaching pressure? A case study of the Diana guenon (Cercopithecus diana) and the western red colobus (Procolobus badius) I. Kone and J. Refisch 11. Vulnerability and conservation of the Tai Forest monkeys W. S. McGraw.

Homoplasy and thick enamel in primates

Traditionally, thick enamel has often been used to infer durophagy (i.e., hard nut and seed consumption) in extinct hominins. These inferences are based on the hypothesis that thick enamel is primarily an adaptation to prevent tooth fracture or chipping resulting from high-stress loads produced during the mastication of large hard foods. An alternative view argues that thick enamel may aid in maintaining tooth function in the face of gradual dental wear from grit, phytoliths and acid, which may be found in foods of widely varying hardness. We use estimates of primate dietary abrasiveness and recorded lifespan to test the hypothesis that enamel thickness is selectively responsive to lifetime dental wear resistance. We use data from the literature to relate enamel thickness to measures of dietary abrasiveness, diet profiles, and longevity for 17 primate species and performed linear regression using several combinations of these variables. We found a positive association between lifetime dietary wear and enamel thickness, suggesting that thick molar enamel in primates may have evolved as a means to resist wear apart from selection to resist tooth fracture. Assuming our estimates of lifetime dietary wear are accurate, we caution against ascribing thick enamel solely to the presence of hard-object feeding in paleoanthropological contexts without also considering primate lifespan and other aspects of feeding ecology.

Positional Behavior of Cercopithecus petaurista

I studied the positional behavior and habitat use of Cercopithecus petaurista, the lesser spot-nosed monkey, in the Ivory Coasts Taï Forest for 15 months. I compare these data with similar information collected on sympatric groups of Cercopithecus diana and C. campbelli in order to examine further the relationships between locomotion, posture, support use, stratal use, body size, diet, activity patterns and foraging behavior. Spot-nosed monkeys are predominantly quadrupedal primates that frequent the top layer of the understory during all maintenance activities. Locomotion is characteristically slow and cautious; travel takes place on branches and boughs while foraging occurs on twigs and branches. Postural behavior of Cercopithecus petaurista reflects their reliance on more ubiquitously distributed, less mobile food items. The relationship between body size, climbing, leaping and support use among Taï guenons is weak; interspecific differences are more likely functions of strata use and overall behavioral characteristics, e.g. crypticity. I also compare the locomotion and support use of Cercopithecus petaurista with that of C. ascanius from Ugandas Kibale Forest (Gebo and Chapman, 1995a) in order to assess the behavioral similarity of members of the same superspecies. Although overall support use is quite similar, the monkeys differ significantly in frequencies of quadrupedism, leaping and climbing. I present possible reasons for and implications of these differences.

Update on the Search for Miss Waldron’s Red Colobus Monkey

Miss Waldron’s red colobus (Procolobus badius waldroni) has a restricted distribution in eastern Ivory Coast and western Ghana. There have been no confirmed sightings of them since 1978 and surveys carried out from 1993 to the present have yet to reveal any living individual. Since the announcement of the monkey’s probable extinction (Oates et al., 2000), new evidence from forest in the extreme southeast of Ivory Coast suggests that a handful of individuals have remained undetected to this point. I discuss the evidence—a tail, a skin and a photograph—and results of accompanying surveys. Additional surveys of the Ehy Forest are required to confirm the presence of Procolobus badius waldroni and an Action Plan for the conservation of all red colobus across the continent is needed to prevent the disappearance of other taxa.Miss Waldron’s red colobus (Procolobus badius waldroni) has a restricted distribution in eastern Ivory Coast and western Ghana. There have been no confirmed sightings of them since 1978 and surveys carried out from 1993 to the present have yet to reveal any living individual. Since the announcement of the monkey’s probable extinction (Oates et al., 2000), new evidence from forest in the extreme southeast of Ivory Coast suggests that a handful of individuals have remained undetected to this point. I discuss the evidence—a tail, a skin and a photograph—and results of accompanying surveys. Additional surveys of the Ehy Forest are required to confirm the presence of Procolobus badius waldroni and an Action Plan for the conservation of all red colobus across the continent is needed to prevent the disappearance of other taxa.

Brief communication: Enamel thickness and durophagy in mangabeys revisited.

The documentation of enamel thickness variation across primates is important because enamel thickness has both taxonomic and functional relevance. The Old World monkeys commonly referred to as mangabeys have figured prominently in investigations of feeding ecology and enamel thickness. In this article, we report enamel thickness values for four mangabey taxa (Cercocebus atys, Cercocebus torquatus, Lophocebus aterrimus, and Lophocebus albigena), offer revised interpretation of the significance of thick enamel in papionin evolution, and place our new data in a broader comparative framework. Our data indicate that all mangabeys have thick enamel and that the values obtained for Cercocebus and Lophocebus equal or exceed those published for most extant non-human primates. In addition, new field data combined with a current reading of the dietary literature indicate that hard foods make up a portion of the diet of every mangabey species sampled to date. Clarification on the relationship between diet and enamel thickness among mangabeys is important not only because of recognition that mangabeys are not a natural group but also because of recent arguments that explain thick enamel as an evolved response to the seasonal consumption of hard foods.

Material property variation of mandibular symphyseal bone in colobine monkeys

The anterior mandibular corpus of anthropoid primates is routinely subjected to masticatory loads that result in relatively high local levels of stress and strain. While structural morphological responses to these loads have been extensively explored, relatively little is known about material property variation in mandibular bone of nonhuman primates. Consequently, the role of regional and local variation in bone stiffness in conditioning stress and strain gradients is poorly understood. We sampled elastic modulus variation in the bone of the anterior mandibular corpus in two species (N = 3 each) of sympatric colobine monkeys, Procolobus badius and Colobus polykomos. These monkeys were chosen for comparison owing to their distinctive dietary regimens, as P. badius rarely includes hard objects in its diet while C. polykomos habitually processes obdurate items during feeding. Elastic modulus is determined through bone hardness data obtained via microindentation, which enables the description of stiffness variation on sub‐millimeter scales. Labial bone stiffness exceeds that of lingual bone in the sample overall. Female mandibular bone is generally stiffer than that found in males, and overall Procolobus mandibular bone is stiffer than that in Colobus. These results, interpreted collectively, suggest that the material response to elevated masticatory stress is increased compliance of the affected bone. J. Morphol., 2009.