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The morphology and wear of the lingual notch in macaques and langurs.

Although interspecific differences in the relative development of molar occlusal features have been documented for most primate species, wear-related changes in these features are only beginning to be understood. In the present study, the notch between the metaconid and entoconid of M2 was examined in a skeletal sample of Macaca fascicularis, presbytis cristatus, and Presbytis rubicunda from the Museum of Comparative Zoology at Harvard University. Photogrammetric analyses yielded X, Y, and Z coordinates that were used to compute three angular measurements and two length measurements at the lingual notch. Statistical analyses of measurements from unworn dentitions indicated that the mesial slope of the entoconid is steeper than the distal slope of the metaconid in all three species. In addition, interspecific comparisons demonstrated that P. cristatus has the steepest lingual notch bordered by the largest molar shearing crests, while P. rubicunda and M. fascicularis have molar shearing crests of similar size, with P. rubicunda merely having higher cusps that are closer together than those of M. fascicularis. Finally, P. cristatus shows a smaller decrease in lingual cusp relief with wear than M. fascicularis, while P. cristatus and P. rubicunda probably show similar decreases in lingual cusp relief with wear. The differences in wear-related changes in lingual cusp relief between M. fascicularis and P. cristatus may well be related to the frugivorous/folivorous dietary differences between these species.

Animals↗

Ontogenetic and interspecific organ weight allometry in Old World monkeys.

The importance of allometry as an analytic tool is well recognized in the literature of primate morphology. However, a number of recent studies have illustrated how interpretive difficulties can arise when researchers confound different types of allometric data. Such confusion is due less to carelessness than to uncertainty about how different types of allometry are related. The present study examines the relationship between two types--ontogenetic and interspecific allometry--in the case of organ weight scaling in six species of Old World monkeys. Accepting the interpretation of interspecific allometry as a reflection of functional scaling constraints, the results of this analysis indicate how ontogenetic patterns have been modified in different-sized species to maintain compliance with these constraints. Specifically, for the heart and lungs it appears that vertical transpositions of individual species' ontogenies are dictated by isometric interspecific allometry, while in the case of the kidneys and liver, the relation of negative allometry across species entails alteration of the relative growth coefficients of the individual species. While these conclusions can at present only be applied to organ weight scaling, the approach of examining interspecific patterns in light of developmental differences between species should prove very helpful in our efforts to understand the phenomena of size and scaling.

Aging↗

Distribution of enamel on the incisors of Old World monkeys.

Longitudinal ground sections of 29 Old World monkey central lower incisors were studied histologically and metrically. Labiolingual incisor width tended to scale isometrically with body weight but with important deviations in relative incisor size, which appeared to be correlated with diet in accord with work by Hylander. Lower incisors of the predominantly folivorous colobine monkeys had a substantial layer of enamel on both lingual and labial aspects and consequently had blunt incisal edges. These teeth in both cercopithecins and papionins, which are omnivorous or frugivorous, had little or no enamel on the lingual aspect, resulting in sharp incisal edges. It is suggested that colobine incisors are used mainly in gripping and tearing leaves, whereas cercopithecine incisors are better adapted to cutting and scraping. Crown height showed a positive allometric relationship with overall incisor height, so that the tall incisors of papionins, especially Papio and Mandrillus, were more hypsodont than the shorter incisors of colobines and cercopithecins. This appears to be related to differences in the rates of incisor wear between the groups.

Animals↗

Nut-like oil seeds: food for monkeys, chimpanzees, humans, and probably ape-men.

The hypothetical hyperrobust australopithecine gnathic nutcracker adaptation is reexamined in light of ecobotanical information on edible wild nuts provided by the flora of tropical and subtropical Africa. The nut producing species are tree-forms. Those of the forest region do not as a rule produce fruits with edible mesocarps. In contrast, the woodland savanna species (particularly in the Zambezian region) characteristically provide an important whole fruit, i.e., a nutritious mesocarp in addition to edible oil-rich nut seeds. These fruits drop from the tree before they are fully mature and go through the final ripening phase on the ground. They are important seasonal foods for a variety of vertebrates, including primates, elephants, and antelope. Altogether the nuts exhibit a broad range of toughness values, measured here as strength under compression. The woodland nuts are not as tough (177-934 kg force, breaking load) as those of the tropical forest (192-1,673 kg force). The seed-predators of the woodland species include squirrels, baboons, warthogs, and parrots. Paleoecological analyses indicate that it was the woodland nuts that were probably available to Australopithecus boisei and A. robustus. Preliminary estimates of adult male gnathic nut-cracking capabilities suggest that A. boisei could have orally cracked a significant portion of the woodland nuts. In spite of this, ecobotanical data indicate that we can probably reject the hypothesis that these hominids were year-round gnathic nut-cracking specialists. Both the indirect and direct evidence support this conclusion.

Africa↗

Seed-eating by West African cercopithecines, with reference to the possible evolution of bilophodont molars.

Data on tooth use in eating fruits and seeds were collected on 12 West African monkeys representing five species of cercopithecines, Cercopithecus aethiops, Cercopithecus campbelli, Cercocebus atys, Erythrocebus patas, and Papio papio. Field observations and information in the published literature were used to select fruits for captive feeding trials. A total of over 27,000 tests were conducted to determine the preferences of the monkeys for 78 fruits. Data were collected on the selection of fruit parts, as well as tooth use, in eating 88 fruit species. All five monkey species had remarkably similar fruit part preferences. Seeds were eaten in 86% of the tested fruits and represent the fruit part most frequently eaten by all the tested monkeys. With few exceptions, molars were used to puncture and crush the seeds. It is suggested that the development of bilophodont molars might be an adaptation by Old World monkeys to seed predation. Given their large size relative to other arboreal frugivores, seed predation could provide a dietary niche for Old World monkeys.

Adaptation, Physiological↗

Absence of strongly kin-preferential behavior by adult female sooty mangabeys (Cercocebus atys).

The objective of this study was to test the hypothesis that kin-preferential behavior would be present in sooty mangabeys (Cercocebus atys), a species taxonomically close to baboons and macaques. The affiliative behavior of the adult female members of a large captive group of these mangabeys was analyzed to test this prediction. These females groomed, were approached by, were in proximity to, and were in contact with their kin significantly more than expected, but only when all kin were included in the analysis. Removal of only the mother-infant (less than 1 year) dyadic interactions removed all significant kin effects. Kin-preferential behavior was also absent in affiliative interactions among the adult females as a class. Affiliation between mothers and offspring significantly exceeded that for all other kinship categories (such as siblings, etc.), and affiliation with kin other than offspring did not differ from that with nonkin adults. In their interactions with nonkin, the adult females showed some preference (duration of grooming) for those adult females of similar age, an effect predicted by the intensity of interaction among members of the same age cohort during development. These similar-aged females may also be paternally related. In comparing these results with the existing literature on kin-preferential behavior, two conclusions may be reached: (1) age and degree of kinship are primary factors that must be considered in order to avoid the existing overgeneralization of the importance of kinship in primate social organization, and (2) the role and importance of affiliative behavior in kin-selection theory should perhaps be re-examined in light of questions raised by this study.

Animals↗

Functional assessment of subfamily variation in maxillomandibular morphology among Old World monkeys.

Among Old World monkeys, subfamily variation in maxillomandibular form is commonly attributed to divergent dietary and social behaviors. However, our knowledge of any musculoskeletal adaptations for gape in cercopithecines, and folivory in colobines, is incomplete. Such data are requisite to a more informed perspective on the evolutionary morphology of these taxa. Structural analyses of gape and biomechanical efficiency were applied to a representative sample of adult cercopithecids. Factors pertaining to the biomechanical scaling of cranial structures were evaluated with least-squares bivariate regression techniques. To assess subfamily differences in masticatory efficiency, analyses of covariance were made between relevant factors. Cercopithecines achieve increased gape and relative canine size mainly with strong positive allometry of the facial skull, combined with a larger gonial angle. Colobines possess a relatively long masseter lever arm and short facial skull, as well as an enlargened masseter-medial pterygoid complex. Subfamily differences in temporalis lever arm scaling are negligible. Biomechanical comparisons within and between subfamilies suggest that the mechanical advantage of the temporalis is relatively greater than that of the masseter, while the mechanical advantage of both muscles increases with face length. Evidence is presented to stress the need for adequate consideration of the dependent variable in allometric investigations of skull form.

Analysis of Variance↗

Comparison of muscle weight and force ratios in New and Old World monkeys.

Thin mandibles and small incisors found in New World monkeys as compared with Old World monkeys suggest that there may be differences in craniofacial loading patterns between these two groups, particularly in levels of mandibular corpus twisting (Hylander, 1975, 1979a; Eaglen, 1984; Bouvier, 1986a,b). This study examined the hypothesis that changes in the relative force contributions of the masticatory muscles were responsible for lowering torsion on the mandibular corpus in New World monkeys. Muscle weight and physiological cross-sections were compared using data from the literature (Schumacher, 1960: Turnbull, 1970; Cachel, 1979) as well as new data on adult male Cebus apella and Macaca mulatta. Both age and sex had an effect on muscle ratios. Mixed samples such as those used by Schumacher and Turnbull probably are not appropriate for drawing conclusions concerning species or group differences in muscle ratios. In addition, biomechanical conclusions based on muscle weight ratios alone to estimate muscle force may be misleading because fiber length inversely affects the amount of force a muscle can exert. A comparison of ratios based on physiological cross-section as an estimator of muscle force in New and Old World monkeys does not support the hypothesis that alterations in force contribution by individual masticatory muscles are responsible for minimizing mandibular corpus twisting in New World monkeys. Therefore, if twisting has been minimized in New World monkeys as suggested by their thin corpora, other changes in the craniofacial musculoskeletal complex, such as different muscle recruitment or pinnation patterns, may be responsible.

Animals↗

Aging and reproductive performance in langur monkeys (Presbytis entellus).

Analysis of data on age and reproductive performance in 16 captive female Hanuman langurs (Presbytis entellus) shows that adult females under the age of 6 years and over the age of 19 years had the lowest rates of reproductive success as measured by the production of viable young that survived beyond 30 days of age. With the possible exception of one female, females over the age of 19 years experienced a variety of idiosyncratic reproductive problems but did not experience a true menopause as defined by total cessation of menstrual cycles. There was no support for the hypothesis that langur monkey females have evolved a lengthy postreproductive period.

Aging↗

Kinematics of the cercopithecine foot on arboreal and terrestrial substrates with implications for the interpretation of hominid terrestrial adaptations.

The stereotyped characterizations of quadrupedal foot postures were tested by examining the kinematics of the cercopithecine foot on arboreal and terrestrial supports. Strictly arboreal species were compared with semi-terrestrial species for Cercopithecus, Cercocebus, Lophocebus, and Papio, in semi-natural or experimental settings. Results indicate that the kinematics of the cercopithecine arboreal quadruped differ in degree from stereotypical expectations for an arboreal quadruped. The relatively extended, adducted limb movements of the cercopithecines and the emphasis on the central digit as the functional axis of the foot suggest convergence with terrestrial mammalian cursors, and differ from the platyrrhine or colobine arboreal quadruped. The characteristics of the quadrupedal terrestrial primate foot contrast with the very unique pattern seen in the hominid foot. These contrasts provide a new perspective from which to interpret the hominid adaptation, in which the functional axis has remained fixed between the first and second digits. This pattern differs from virtually all other terrestrial mammals. The influence of bipedalism on this functional pattern is examined.

Animals↗

Facial anatomy of Victoriapithecus and its relevance to the ancestral cranial morphology of Old World monkeys and apes.

Recently discovered craniofacial fossils of the middle Miocene cercopithecoid Victoriapithecus are described. The frontal, zygomatic, maxilla, and premaxilla anatomy differ from the previously proposed colobine-like ancestral cercopithecoid morphotype in several significant respects. This morphotype was based on the assumption that features held in common by subordinate hominoid and cercopithecoid morphotypes (Colobinae and Hylobatidae) are primitive for Old World monkeys. Cranial similarities between Victoriapithecus, which represents the sister-group of both colobine and cercopithecine monkeys, and the shorter-snouted Cercopithecinae (Macaca and Cercopithecus) indicate that the last common ancestor of Old World monkeys possessed the following features: a narrow interorbital septum, moderately long snout, moderately long and anteriorly tapering premaxilla, large procumbent upper central incisors set anterior to and with longer roots than lateral incisors, moderately tall face below the orbits, teardrop-shaped nasal aperture of low height and moderate width, and probably long and narrow nasal bones. The Victoriapithecus cranium is also characterized by features not present in modern cercopithecids. These include a deep malar region of the zygomatic and the presence of a frontal trigon due to the occurrence of temporal lines that merge with supraorbital costae close to the midline of each orbit and converge anterior to bregma. These features are interpreted as primitive retentions from the basal catarrhine condition as indicated by the occurrence of these features among primitive catarrhines (Aegyptopithecus) and Miocene hominoids (Afropithecus).

Animals↗

Variability at the carpometacarpal and midcarpal joints involving the fourth metacarpal, hamate, and lunate in catarrhini.

Intraspecies variability is investigated in two regions of the wrist, for the purpose of determining whether patterns may be discerned in the variability that may be compared in the functional and phylogenetic analysis of living and fossil catarrhines. In the midcarpal joint region, two lunatohamate configuration patterns are found, and at the fourth carpometacarpal joint four types of configuration are identified. These two sites previously were reported to show almost continuous variability in humans, thus precluding comparison with other species. The different types of configuration in our study are delineated on the basis of their relation to differences in joint function. At the lunatohamate site there is a strong tendency in each species examined for one type to dominate in frequency. At the fourth carpometacarpal joint there is a tendency for one type or for two related types to predominate in each species. The chimpanzee sample exhibits the least variability of all species studied in joint configuration at the two sites. Australopithecus afarensis has a combination of joint types in these regions likely to be found today in only a small percentage of living Hominoidea. We conclude that patterns may be discerned at some joints in what was formerly considered to be a continuum of variation. Since these patterns (joint types) differ in their relative frequencies among living species, the frequency differences may be useful as a guide to the reconstruction of phylogenetic relationships and of potential wrist functions in fossil species.

Animals↗

Incisor microwear of Sumatran anthropoid primates.

Several studies have suggested that incisor microwear reflects diet and feeding adaptations of anthropoids. However, such studies have been largely qualitative, and interpretations have relied on anecdotal references to diet and tooth use reported in the socioecology literature. The current study relates incisor microwear in four anthropoid primates to specific ingestive behaviors and food types. Central incisor casts of wild-shot museum specimens of Hylobates lar, Macaca fascicularis, Pongo pygmaeus, and Presbytis thomasi were examined by scanning electron microscopy, and analyzed using a semiautomated image analysis procedure. Microwear patterns were used to generate predictions regarding diet and anterior tooth use. These predictions were evaluated using data collected during a 1 year study of feeding behavior of these same taxa in the wild (Ungar, 1992, 1994a,b). Results suggest that (1) enamel prism relief is associated with the effectiveness of etching reagents in foods, (2) dental calculus buildup results from a lack of incisor use and perhaps the ingestion of sugar-rich foods, (3) striation density varies with degree of anterior tooth use in the ingestion of abrasive food items, (4) striation breadth is proposed to relate to the ratio of exogenous grit to phytoliths consumed; and (5) preferred striation orientation indicates the direction that food items are pulled across the incisors during ingestion. It is concluded that incisor microwear studies can contribute to the understanding of diets and feeding behaviors of extinct primates.

Analysis of Variance↗

Ontogenetic correlates of diet in anthropoid primates.

This study assesses ontogenetic correlates of diet in anthropoid primates. Associations between body weight growth, adult size, and diet are evaluated for a sample of 42 primate species, of which 8 are classifiable as "folivores." The hypothesis that folivores show a pattern of growth that differs from "nonfolivores" is tested. Ontogenetic variation is summarized through use of parametric and nonparametric regression analysis. Several analytical techniques, including broad interspecific and detailed comparisons among species of similar adult size, are applied. This investigation indicates a clear association between body weight ontogeny and diet: folivorous species grow more rapidly over a shorter duration than comprably sized nonfolivorus species. A positive correlation between adult size and diet is not unambiguously established in this sample. A threshold (at around 1 kg) below which insectivory is very common may adequately characterize the association between adult size and diet in anthropoid primates. Above this threshold, adult size does not appear to covary predictably with diet. Evolutionary correlates of the ontogenetic pattern seen in folivores may include a variety of factors. The distinctive pattern of development in folivores may relate to the profile of ecological and social risks that these species face. Morphophysiological advantages to rapid growth may relate to a need for accelerated alimentary (dental and gut) development. The implications of ontogenetic variation in folivores are discussed.

Age Factors↗

Patterns of ingestive behavior and anterior tooth use differences in sympatric anthropoid primates.

Little research has been directed towards the examination of ingestive behaviors in wild primates. This paper describes a naturalistic study of anterior tooth use in four sympatric anthropoid species: Hylobates lar, Macaca fascicularis, Pongo pygmaeus, and Presbytis thomasi. Instantaneous group scan data were collected during nearly 1,800 hours of observation between August 1990 and July 1991 at the Ketambe Research Station in the Gunung Leuser National Park, Sumatra, Indonesia. Ingestive behaviors are documented for specific food items and compared among the primate taxa. Results indicate significant differences among the species in preferred methods of food ingestion. These differences are related in part to dietary differences, and in part to other aspects of each primate's biology and ecology.

Animals↗

Dust accumulation in the canopy: a potential cause of dental microwear in primates.

Dental microwear researchers consider exogenous grit or dust to be an important cause of microscopic wear on primate teeth. No study to date has examined the accumulation of such abrasives on foods eaten by primates in the forest. This investigation introduces a method to collect dust at various heights in the canopy. Results from dust collection studies conducted at the primate research stations at Ketambe in Indonesia, and Hacienda La Pacifica in Costa Rica indicate that 1) grit collects throughout the canopy in both open country and tropical rain forest environments; and 2) the sizes and concentrations of dust particles accumulated over a fixed period of time differ depending on site location and season of investigation. These results may hold important implications for the interpretation of microwear on primate teeth.

Alouatta↗

Anatomy of the nasal cavity and paranasal sinuses in Aegyptopithecus and early Miocene African catarrhines.

Neontological comparisons suggest that paranasal sinus anatomy is diagnostic of several catarrhine clades such as Cercopithecoidea, Hominoidea, Homininae, and Ponginae. However, while the loss of sinuses in cercopithecoids is generally recognized as a derived condition, determining the polarity of character-state changes within noncercopithecoid catarrhines requires knowledge of the primitive catarrhine condition. To address this problem, the paranasal sinus anatomy of Aegyptopithecus and several early Miocene catarrhines was investigated. Two partial facial skeletons of Aegyptopithecus were subjected to computed tomography in order to reveal their internal anatomy. These data were compared with facial and palatal specimens of Proconsul, Limnopithecus, Dendropithecus, Rangwapithecus, and Kalepithecus in the National Museums of Kenya in Nairobi, and to wet and dry specimens of living taxa. Results confirm that cercopithecoid paranasal anatomy is derived, and reveal that the sinus anatomy of stem catarrhines included a hominoid-like maxillary sinus as well as an ethmofrontal system like that of hominines. Accordingly, these two features do not constitute evidence for the hominoid, hominid, or hominine status of any fossil species. Conversely, the absence of the ethmofrontal sinus system in Sivapithecus and Pongo is synapomorphic. In addition, features of the nasal cavity of Limnopithecus and Kalepithecus support previous suggestions that these taxa are stem catarrhines rather than hominoids.

Africa↗