Ischial tuberosities and ischial callosities

Ischial callosities are specialised regions of skin and subdermal tissue in the form of fibro-fatty cushions with a tough, non-slip surface, firmly bound to the underlying ischial tuberosity. They occur in primates of the families Cercopithecidae and Hylobatidae. Ischial callosity usage helps animals to adopt stable sitting postures on the tops of branches, particularly during feeding, resting and sleeping. Callosity-like areas in pongids are mainly epidermal specialisations and differ in their development and usage from true callosities. They are similar to the para-callosity skin regions in some ground living cercopithecids. The ischial tuberosity in animals with true callosities has a flat surface which flares out from the lower end of the ischial body and which is devoid of muscle attachments. In animals without callosities the ischial tuberosity is not flared, has a curved surface, and acts as a site for muscle attachment. The ischial tuberosity is relatively broader in animals with callosities than in animals without callosities, although the absolute size of the specimens concerned must be taken into account. The combination of qualitative and quantitative features of fossil ischial tuberosities enables the presence or absence of callosities in these forms to be deduced. The recognition of these features in fossil material might be of use in the elucidation of the evolution of one type of primate arboreal adaptation.     

Comparative assessment of the ischial morphology of Victoriapithecus macinnesi

An almost complete primate ischium was recovered from middle Miocene (ca. 15 ma) deposits of Maboko Island (Kenya) in 1987. The specimen shows numerous similarities to the ischial morphology of extant cercopithecids and is attributed to Victoriapithecus macinnesi (Von Koenigswald, 1969), the early Old World monkey best known from Maboko Island. The Victoriapithecus ischium provides the first evidence of early Old World monkey pelvic girdle anatomy. The ischium is characterized by an obliquely oriented and broadly flaring tuberosity, a relatively small acetabulum with little ventrally directed curvature of its caudal portion, a long ischial body and a flange-like ischial spine positioned caudal to the rim of the acetabulum. In these features, Victoriapithecus most closely resembles the vervet monkey, Cercopithecus aethiops. The fossil specimen indicates that Victoriapithecus possessed ischial callosities, a mobile tail and adaptations for (possibly cursorial) quadrupedalism with an adducted posture of the thigh. The occurrence of ischial callosities in Victoriapithecus extends the documented antiquity of this feature in catarrhines by more than 12 million years and shows that the distinctive “sitting-sleeping” adaptations of Old World monkeys (Washburn, 1957) originated prior to the divergence of Colobinae and Cercopithecinae. Differences of developmental sequence and tissue composition indicate that the ischial pads of cercopithecids, hylobatids, and pongids may have arisen independently, through parallel evolution. Contrary to Strasser and Delson (1987), discontinuity of ischial callosities was probably the primitive condition for male cercopithecids.     

Functional aspects of primate pelvic structure: A multivariate approach

This study aims to clarify the relationship of primate bony pelvic structure to locomotor habit. As with most of the postcranial skeleton, the pelvic bones of species within the Ceboidea and the Cercopithecoidea are remarkably similar visually except for variations in size. Yet there are substantial differences in locomotor pattern between the species in these taxa. I performed canonical analyses on a sample of 17 pelvic variables describing 22 primate species of the Ceboidea, the Cercopithecoidea, and the Hominoidea to discover which variables were significant in separating them into groups. In both analyses there was good separation of major taxa and additional separation of groups that differed in locomotor habit. The separation of colobine from cercopithecine monkeys was particularly consistent. In the analysis, including all 22 species, the variables given particular weight by the canonical analysis were the same as those traditionally used by anatomists for the same purpose. Specifically, breadth of the ischial tuberosity (reflecting presence or absence of ischial callosities) separated the Old from the New World monkeys. Breadth of the iliac tuberosity, in which man and to some extent other hominoids differ from other primates, and ilium height, in which man differs from other primates, were significant. Sagittal diameter of the pelvis was also substantially weighted. Having established that the technique would select variables of anatomical significance, the same method was applied to a study of monkeys only where the characteristics that differ between groups are not well established. Breadth of the ischial tuberosity was again important in separating the Ceboidea from the Cercopithecoidea. Discrimination of locomotor groups within these large divisions was brought about mainly by ischial length and the sagittal diameter of the pelvis. In studying these variables and their relationship to size in greater detail, it was found that among cercopithecoid monkeys, the colobines showed relatively lower values than did cercopithecines for both these dimensions. Atelines showed low values for ischial length but high values for the sagittal pelvic diameter. Biomechanical explanations of these observations are suggested.     

Activity budget and diet of patas monkeys in Kala Maloue National Park,Cameroon: A preliminary report

A preliminary study was carried out on the feeding ecology of patas monkeys in the rainy season in Cameroon. Their daily activity rhythm revealed two active peaks. The proportion of time spent on feeding with respect to waking time was 30%. Patas monkeys largely depended on the flowers and buds of herbaceous plants and the larvae of insects for their diet as they ranged widely. Patas monkeys spent more time in feeding and travelled for a longer distance per day than the sympatric primate species, the tantalus monkey. It is considered that these findings reflected the large amount of food requirement due to the large body size, as well as the low density and high degree of dispersal of their food.     

Positional behavior,selective use of habitat substrate and associated non-positional behavior in free-rangingColobus guereza (Rüppel, 1835)

Positional behavior (posture and locomotion), associated social/maintenance behaviors and selective utilization of available habitat structure were recorded for free-rangingColobus guereza occupying a remant montane forest near Limuru, Kenya. This research design focuses on relationships between anatomy, behavior, and the environment and provides a framework for interpreting adaptations inColobus. Quantitative analyses show daily activity patterns. These monkeys spend most of their time in above branch postural activities, primarily in sitting postures. Locomotion accounts for less than one-fifth of positional behavior. In both postural and locomotor sequences, the forelimbs may be placed above the shoulder complex, often in abducted positions. Suspensory positional behavior is rare. Most activity occurs within Zone II, flexible branches of medium diameter and less than 45° in grade. Selective use of the substrate and preferred positional modes are related to variables such as social/maintenance activities, weight distribution of the link system and substrate configuration. These data suggest that, (a) black and white colobus do not exhibit behavior associated with so-called “semibrachiation” and (b) bio-behavioral adaptations involving foraging and feeding strategies may account, in part, for observed morphological differences among Old World monkeys.     

Program of the fifty-sixth annual meeting of the American Association of Physical Anthropologists to be held in New York City Vista International Hotel April 2–5, 1987

Male orangutans (Pongo pygmaeus) weigh about twice as much as females. Observations in northern Sumatra of adults of both sexes (one male, three females) feeding in the same trees reveal the effects of body size dimorphism on feeding behavior: The male tended to use larger branches than the females, and to employ above-branch postures (sitting and standing) with greater frequency. The females employed suspensory under-branch postures more often. When feeding techniques were variable, the male tended to pull in branches to detach food with the mouth, whereas the female plucked more fruit by hand. By controlling for postcranial morphology and habitat structure, these results provide the first rigorous quantitative test of predictions about the effects of body size on primate positional behavior, and raise further questions about sexual diethism in feeding postural behavior of primates of varying absolute size.     

A Comparative Assessment of Hand Preference in Captive Red Howler Monkeys,Alouatta seniculus and Yellow-Breasted Capuchin Monkeys,Sapajus xanthosternos

There are two major theories that attempt to explain hand preference in non-human primates–the ‘task complexity’ theory and the ‘postural origins’ theory. In the present study, we proposed a third hypothesis to explain the evolutionary origin of hand preference in non-human primates, stating that it could have evolved owing to structural and functional adaptations to feeding, which we refer to as the ‘niche structure’ hypothesis. We attempted to explore this hypothesis by comparing hand preference across species that differ in the feeding ecology and niche structure: red howler monkeys, Alouatta seniculus and yellow-breasted capuchin monkeys, Sapajus xanthosternos. The red howler monkeys used the mouth to obtain food more frequently than the yellow-breasted capuchin monkeys. The red howler monkeys almost never reached for food presented on the opposite side of a wire mesh or inside a portable container, whereas the yellow-breasted capuchin monkeys reached for food presented in all four spatial arrangements (scattered, on the opposite side of a wire mesh, inside a suspended container, and inside a portable container). In contrast to the red howler monkeys that almost never acquired bipedal and clinging posture, the yellow-breasted capuchin monkeys acquired all five body postures (sitting, bipedal, tripedal, clinging, and hanging). Although there was no difference between the proportion of the red howler monkeys and the yellow-breasted capuchin monkeys that preferentially used one hand, the yellow-breasted capuchin monkeys exhibited an overall weaker hand preference than the red howler monkeys. Differences in hand preference diminished with the increasing complexity of the reaching-for-food tasks, i.e., the relatively more complex tasks were perceived as equally complex by both the red howler monkeys and the yellow-breasted capuchin monkeys. These findings suggest that species-specific differences in feeding ecology and niche structure can influence the perception of the complexity of the task and, consequently, hand preference.     

Use of forest canopy by European red squirrelsSciurus vulgaris in Northern Greece: claws and the small branch niche

Moving and standing in trees impose multiple problems to arboreal mammals. Among them, the major ones are the negotiation of slender terminal branches and of large vertical supports. Both microhabitats are important as they have been linked alternatively to the evolutionary loss of claws in early primates. Therefore, rates of use of these different supports by claw-bearing arboreal mammals may offer insights to their actual significance in the adaptive evolution of early primates. In this context, canopy, tree crown, branch size, inclination, and texture use were recorded on four adult free ranging European red squirrelsSciurus vulgaris Linnaeus, 1758 in a mixed coniferous forest in northern Greece.S. vulgaris was mainly arboreal, exploiting the terminal branch zone, using frequently oblique and intermediately textured supports<5 cm and moderately large vertical branches. Furthermore, comparative data from other sciurid species and clawed primates showed positive correlations of small and horizontal support use, and negative ones of vertical support use to body mass. These findings show that keeled functional claws do not impede habitual use of slender branches and may not facilitate efficient climbing on large vertical trunks. These observations partly question the association between habitual use of the small branch niche and primate adaptations and lend support to alternative hypotheses, underscoring the importance of inquiring for more complex mechanisms that lead to the evolution of the unique set of primate morphological adaptations.     

The behavioral ecology of three Indian Ocean surgeonfishes (Acanthurus lineatus,A. leucosternon and Zebrasoma scopas): their feeding strategies,and social and mating systems

Synopsis The relationship between the morphology, feeding strategies and social and mating systems of three surgeonfishes was investigated. Adults of each defend feeding territories, intra-and interspecifically. The largest species, because of its morphological limitation, relies on food that has to be defended against many other species. It forms large colonies in which fishes singly defend small territories containing high standing crop algal mats. Colony formation is a mechanism by which the efficiency and effectiveness of interspecific territory defense is increased. The smallest species, because of its morphological adaptations, is able to rely most on food that other species cannot efficiently exploit. It forms pairs that defend large territories containing a thin algal mat. It is restricted to the poorest quality habitat by the aggressive activities of more dominant species. The third species, which also forms pairs, has an intermediate feeding strategy. The local coexistence of these three and other surgeonfishes results from a combination of (i) their partitioning both habitat and food resources, and (ii) the populations of two of the most dominant species apparently being below the carrying capacity. Territoriality and the absence of parental care facilitates pair formation in surgeonfishes. Permanently territorial species usually form pairs. The colonial species does not form pairs because the colonial habit facilitates interference of males in each other's spawnings.     

The Narrow Niche hypothesis: gray squirrels shed new light on primate origins

Current hypotheses for primate origins propose that nails and primate-like grasping hands and feet were important early adaptations for feeding in fine branches. Comparative research in this area has focused on instances of convergence in extant animals, showing that species with primate-like morphology feed predominantly from terminal branches. Little has been done to test whether animals without primate-like morphology engage in similar behavior. We tested the fine-branch niche hypothesis for primate origins by observing branch use in Eastern gray squirrels, Sciurus carolinensis, a species lacking primate grasping adaptations that has been understudied in the context of primate origins. We hypothesized that because gray squirrels lack primate-like grasping adaptations, they would avoid feeding and foraging in terminal branches. Instantaneous focal animal sampling was used to examine the locomotor and postural behaviors used while feeding and foraging. Our results demonstrate habitual and effective usage of terminal branches by gray squirrels while feeding and foraging, primarily on tree seeds (e.g., oak, maple, and elm). Discriminant function analysis indicates that gray squirrels feed and forage like primates, unlike some other tree squirrel species. Given the absence of primate-like features in gray squirrels, we suggest that although selection for fine-branch foraging may be a necessary condition for primate origins, it is not sufficient. We propose an alternative model of primate origins. The Narrow Niche hypothesis suggests that the primate morphological suite evolved not only from selection pressure for fine branch use, but also from a lack of engagement in other activities.     

FACTORS GOVERNING FEEDING RATE, FOOD REQUIREMENT AND BROOD SIZE OF NESTLING GREAT TITS PARUS MAJOR

SUMMARY Observations were made on feeding rates and food-consumption of nestling Great Tits Parus major mainly in Larch plantations at lake Yamanaka, Japan. Feeding frequencies were recorded by an automatic recorder. There were marked differences between early and late broods; the feeding frequencies were twice as great in early than in late broods of the same size. No clear tendency was observed in the variations of feeding frequencies in relation to brood size. There was, however, a clear inverse relationship between the frequencies and the average size of food brought to the nests. The males' share in terms of feeding frequencies is described. These figures, however, did not follow the males' contribution in terms of weight of food, which was nearly always higher than the females'. It is pointed out that feeding frequencies are far too variable to be used as a true index of food consumption by nestlings, and are not reliable. Attempts were made to measure the weight of food; the method is described. The average weight of food brought by males was lighter in early than in later broods. The total weight of food was estimated. The trend of daily food consumption per chick was similar to that of the chick's growth curve. It was found that up to about the tenth day of the nestling period daily food-intake per chick increased linearly as body weight increased. At some nests, rate of defaecation was observed. This was at first low, but it increased steeply on the third day, with a steady increase thereafter. By comparing the rates of food intake, faeces output, and weight increment of a chick, it was found that only 20–30% of digested matter (the difference between food-intake and faeces-output was used up daily (for body temperature regulation various external effort, etc.). The factors responsible for this high efficiency of growth in nestlings are discussed. There was a clear inverse relationship between the total weight of food brought per chick per day and the brood size. This is largely because the heat-loss is greater in small than in large broods, so that a chick from a small brood in fact needs more energy to maintain its body temperature after a certain age than one from a large brood. This is discussed in detail. Factors which caused variations in size of food are discussed in relation to feeding frequencies. It is pointed out that, because of the inverse relationship between energy requirement by each chick and brood size, the total food requirement by a brood as a whole did not vary directly in proportion to the brood size. An estimation showed that a b/3 still required about 75% of the total food required by a b/8. A smaller brood is less advantageous than expected to parents feeding nestlings when they encounter adverse conditions, e.g. food shortage in the habitat, or a lack of help by their mates, etc. On the other hand, it is suggested that once they have left the nest, the food-demand by a brood of fledglings the parents have to feed, so that, in the fledging period, in times of food shortage it would certainly be advantageous to have fewer young. It is suggested that, although fledglings may consume three to four times as much food as nestlings, the parents, in providing this food, would not work proportionately harder, since the parents' efficiency of providing food could be higher in feeding the fledglings, which always follow the parents as they are hunting, than in feeding the nestlings to which food has to be brought. On this basis, the adaptive significance of the length of the nestling period in nidicolous species is discussed in relation to clutch size, brood size and food requirement.     

Life-history traits of two Mediterranean lizard populations: a possible example of countergradient covariation

The trade-off between clutch and offspring size, which is a central topic in life-history research, is shaped by natural selection to maximize the number of surviving offspring, but it also depends on the resources available for reproduction. Conspecific populations living in different environments may differ in adult body size, clutch mass, clutch size, offspring size, and/or post-natal growth rates, due either to phenotypic plasticity or to local adaptation. Here, we compare these traits and their relationships between two populations of the lizard Psammodromus algirus separated by a 600-m altitudinal gradient. We used a common garden design to control incubation temperature and food availability, with two different feeding treatments. Females were larger at the high-elevation site. Although SVL-adjusted clutch mass did not differ between populations, high-elevation females laid more but smaller eggs than low-elevation ones. Hatchlings were larger at lower elevation. Our common garden experiment revealed that low-elevation hatchlings grew faster than high-elevation hatchlings under both feeding treatments. However, higher food availability at higher altitude allows high-elevation lizards to grow faster and attain larger adult sizes, especially in the case of females. The two key adaptations of low-elevation lizards, large eggs and hatchlings and the ability to grow rapidly after hatching, are likely to enhance survival in low-productivity Mediterranean lowlands. Our data support the hypothesis that the reproductive strategies of these populations provide an example of countergradient variation, because the genotypes that encode for fast growth and large body size occurred in low food availability habitats where juveniles grew slowly and attained small adult sizes.     

Maximum ingested food size in captive strepsirrhine primates: Scaling and the effects of diet

Little is known about ingested food size (V_b) in primates, even though this variable has potentially important effects on food intake and processing. This study provides the first data on V_b in strepsirrhine primates using a captive sample of 17 species. These data can be used for generating and testing models of feeding energetics. Strepsirrhines are of interest because they are hypometabolic and chewing rate and daily feeding time do not show a significant scaling relationship with body size. Using melon, carrot, and sweet potato we found that maximum V_b scales isometrically with body mass and mandible length. Low dietary quality in larger strepsirrhines might explain why V_b increases with body size at a greater rate than does resting metabolic rate. Relative to body size, V_b is large in frugivores but small in folivores; furthermore scaling slopes are higher in frugivores than in folivores. A gross estimate of dietary quality explains much of the variation in V_b that is not explained by body size. Gape adaptations might favor habitually large bites for frugivores and small ones for folivores. More data are required for several feeding variables and for wild populations. Am J Phys Anthropol 142:625–635, 2010. © 2010 Wiley‐Liss, Inc.     

Time budgets of grassland herbivores: body size similarities

Summary The summer (May–September) time budgets of 14 generalist herbivore species living in the same grassland environment are presented in terms of various component activities (e.g., walking, feeding, resting, etc.). All the species exhibit a decrease in activity as average daily air temperature increases. Greater body size and variety of habitats used by a species lead to increased time spent active. Use of a greater variety of habitats may increase activity time because different habitats provide suitable thermal conditions for activity at different times of the day. Body size affects sn herbivore's thermal balance through metabolism, body surface area and thermal inertia. The time spent feeding, exclusive of time spent searching for foods, is less for large than small herbivores. This may arise because large species must spend more time walking in the search for food to satisfy their energy requirements. The observed feeding time differences for species composing a common trophic level in a single environment may help to explain their diet choice because feeding time constrains the variety of foods an herbivore can select. Diet differences, in turn, can explain the potential competition for food if food is in short supply.     

Use of the forest canopy by the agile gibbon

Brachiation was the main form of locomotion and took place along branches and boughs in the upper canopy. The posture during feeding depended on the substrate, with the gibbons sitting on branches but hanging in twigs. During foraging the gibbons hung more, even on branches, but as only short stops were made this is probably to increase mobility. Feeding on fruit took place mainly in the middle canopy on small trees; feeding on leaves and foraging occurred evenly throughout the middle and upper canopies. Gibbons are light mobile animals with a rapid and direct form of locomotion that enables them to exploit dispersed food sources and to defend a large territory.     

Social rank and body size as determinants of positional behavior inPan troglodytes

A yearlong study of the positional behavior ofPan troglodytes at the Mahale Mountains National Park yielded 571 hr of observation.Cant (1987) articulated four predictions concerning the relationship between body weight/branch diameter and positional behavior based on the classic suspensory-ape paradigm. He noted that only two were supported by orangutan data. Three of these four hypothesis were not supported by chimpanzee data, as follows: there was no significant difference between the three largest males and the four smallest males in (1) branch diameters (5.8 cm vs 5.2 cm) nor (2) in the percentage of arm-hanging (13.6% vs 12.1%); and (3) large males did not arm-hang significantly more often than small males in any of three support diameter categories. The fourth hypothesis, that arm-hanging should be more common among smaller branches, was supported: arm-hanging as a percentage of all posture rose from 2.5% to 8.3% to 24% as stratum size decreased from >10 cm to <3 cm. The possibility that the first three hypotheses failed because of confounding effects of a correlation between body size and social rank was examined. Multiple regressions were done on 6600 2-min instantaneous observations on focal individuals. With social rank effects factored out, larger individuals preferentially utilized smaller, rather than larger supports. When positional mode frequencies were compared between large and small males matched for social rank, large males exhibited a lower frequency of arm-hanging than small males. An unexpected result was that social rank more consistently predicted branch diameter choice than body size. The most profound trend was for high ranking males to use larger supports, even though they spent more time in the terminal branches. These results suggest that (1) suspensory behavior is functionally related to small branch diameters; (2) chimpanzees do not prefer smaller branches, rather they are forced into them by food choice limitations; and (3) social rank more profoundly affects chimpanzee behavior than body weight.     

A Proposal Regarding the Source of Nutrition of Leptocephalus Larvae

The large size, transparency and peculiar dental configuration of leptocephalus larvae can be interpreted as adaptations to a diet of large, transparent and soft food items. It is suggested that gelatinous zooplancton, in particular the discarded house of the Appendicularia or Larvacea, is their main food. Calculations show that an ingestion rate of the order of magnitude 10 houses/ hour is sufficient to sustain an eel larva.     

Maintenance of above-branch balance during primate arboreal quadrupedalism: coordinated use of forearm rotators and tail motion

Animals that live and travel in trees display a variety of morphological and behavioral adaptations to help them maintain balance on narrow flexible supports. Among these adaptations are long tails that can be used as counterweights, and freely mobile limbs in order to reach discontinuous supports. Here we describe two additional ways in which these features can contribute to balance during arboreal locomotion. Electromyographic (EMG) recordings of the forearm rotators pronator quadratus and supinator during over-ground and above-branch quadrupedal locomotion in five species of Old World monkeys revealed their contribution to shifting the weight of the body to help change the direction of travel and maintain balance on a branch. In addition, we observed a coordinated mechanism consisting of a sweeping tail rotation toward the direction of imbalance, to impart an angular momentum to the body that assists in the restoration of balance. While all five primate species utilized forearm rotators to shift their bodies toward one side or the other during quadrupedal walking along a branch, the tail-whip mechanism was most frequently used by the largest and most terrestrial species. We suggest that their large size and/or terrestrial habits have made them less adept at arboreal locomotion, and therefore most likely to utilize auxiliary balancing mechanisms. The usefulness of a long tail as a balancing aid during arboreal locomotion highlights the puzzling nature of the evolutionary loss of a tail in the ape and human lineage.     

A biomechanical analysis of mandibular scaling in old world monkeys

Scaling of mandibular dimensions in male Old World monkeys was investigated. Mandibular condyle length, width, and area were regressed separately against body mass and mandibular length for a total of 14 species of Cercopithecoids. Scaling of mandibular depth and width against both body mass and mandibular length were also investigated. When results of regression analysis using the two different independent variables (body mass and mandibular length) were compared, there were significant pattern differences in scaling of cercopithecines versus colobines. Compared to body mass, male cercopithecines had relatively large mandibles (length, width, and depth) and also relatively large condyles (length, width, and area). However, compared to mandibular length, cercopithecines had relatively transversely thin and shallow mandibles and relatively narrow condyles. It is shown that a “biomechanical” interpretation of mandibular scaling patterns against body mass in Old World monkeys demonstrates only that cercopithecines have prognathic faces, an already well-known and well-documented condition. When the biomechanical effects of prognathic faces are controlled for (by scaling against mandibular length), it is shown that cercopithecines possess special adaptations in condyle length while colobines possess special adaptations in condyle width and mandibular depth and width. These results clearly demonstrate the importance of selecting a relevant reference variable in scaling studies where biomechanical interpretations are attempted.