Atelinae adaptations: behavioral strategies and ecological constraints.

Comparisons between the four genera that make up the Atelinae reveal two distinct behavioral patterns, one in which energy expenditure is minimized (Alouatta) and one in which energy intake is maximized (Lagothrix, Ateles, and Brachyteles). Among the atelins, Lagothrix and Ateles devote over 75% of their annual feeding time to fruit, while Brachyteles devotes between 50% and 67% of their feeding time to leaves. Pronounced seasonality in the Atlantic coastal forest inhabited by Brachyteles may be responsible for its more folivorous diet. Alouatta falls in the body size range of Lagothrix and is much smaller than Ateles and Brachyteles. Nonetheless, Alouatta is more folivorous than sympatric atelins. The atelins also share a rapid, suspensory mode of locomotion that appears to enable them to minimize travel time between widely dispersed fruit sources. Alouatta, by contrast, employs a slower, but more energetically efficient, quadrupedal locomotion. Ranging patterns among the Atelinae are consistent with both diet and locomotor abilities: Atelins travel daily distances up to 5,000 m; Alouatta ranges are much shorter. Further distinctions are evident in Atelinae grouping patterns. Alouatta remains in small cohesive groups that occupy home ranges less than 60 ha in size. Both Lagothrix and Ateles have large groups that fission to reduce the costs of intragroup feeding competition when preferred fruits occur in small patches within much larger community ranges. While greater reliance on low-energy foods such as leaves may release Brachyteles from similar competitive constraints, their tendency toward fluid grouping associations is consistent with the pursuit of a frugivorous diet.     

Tale of tails: parallelism and prehensility

The occurrence of prehensile tails among only five platyrrhine genera--Cebus, Alouatta, Lagothrix, Ateles, and Brachyteles--might be interpreted as evidence that these are a closely related, possibly monophyletic group. In the absence of behavioral data, it is impossible to test whether all possess equivalent biological roles; such would lend credence to the idea that their tails evolved from an homologous, derived character complex. Contrariwise, the tendency for species of Cebus to have "averagely" proportioned or relatively short tails, in contrast to the relatively elongate tails of howlers and other atelines; osteological differences in caudal and sacral morphology; and a lack of ateline-like tail/neocortex correlates in Cebus, all imply that prehensility has evolved twice in parallel: once (homologously) in atelines and again in capuchins.     

Diet and nutrition in wild mongoose lemurs (Eulemur mongoz) and their implications for the evolution of female dominance and small group size in lemurs

Data collected on the feeding behavior, food intake, and chemical analyses of plant foods were used to document seasonal variation in diet and nutrition in Eulemur mongoz in northwestern Madagascar. E. mongoz conforms to the general Eulemur dietary pattern, with a predominantly frugivorous diet supplemented mainly by leaves, flowers, and nectar. Phytochemical analysis revealed high water contents in all the main plant foods; mature fruit and flowers contained the most water-soluble carbohydrates; immature leaves were richest in protein and essential amino acids; the limiting amino acids in all plant foods were methionine and cystine; ash (mineral) content was highest in petioles and mature leaves; crude lipid content was highest in seeds; and crude fiber content was indistinguishable between immature and mature fruit and leaves. High-fiber foods were eaten during both seasons; the wet season diet was dominated by high-energy foods (mature fruit, nectar, and seeds), while the dry season diet contained foods high in energy (mature fruit and flowers) and high in protein (immature leaves) and minerals (mature leaves and petioles). However, nutrient intake did not vary between seasons, implying that nutrient requirements are met throughout the year. These results suggest we draw more conservative conclusions when interpreting dietary variability in the absence of chemical analysis, and also draw into question the idea that nutritional stress is a factor in the timing of reproduction in lemurs and, by extension, is linked to the prevalence of female dominance and small group size in lemurs.     

Seasonal variation in the feeding behavior and diet of Japanese macaques (Macaca fuscata yakui) in lowland forest of Yakushima

The feeding behavior of the southern subspecies of Japanese macaque (Macaca fuscata yakui) was studied over a period of 18 months in warm temperate broad-leaved forest on the island of Yakushima, Japan. Focal animal data were collected for the eight adults in the troop. Over a full annual cycle, 35.0% of foraging on identified foods was on leaves and shoots, 30.2% on fleshy fruit, 13.2% on seeds, and 5.5% on flowers. Invertebrates and other animal matter accounted for 10.3% of foraging and fungi for 4.6%. There was marked seasonal variation in the use of different food categories, and seeds, leaves, fleshy fruit, and animal matter were each predominant at different times of year. There was also evidence of annual cyclicity in patterns of foraging on all major food types. The monkeys spent less time moving and ate a greater variety of foods when feeding on leaves than when feeding on fruit and seeds, or on insects. Time spent foraging was positively correlated with diversity of the diet, but there was no simple relationship between time spent foraging and the predominant food type. This suggests that a wide variety of foods takes longer to harvest and process, irrespective of the food type. The diet of the study troop was flexible and could not be assigned to a simple dietary category, such as frugivorous or folivorous. If these data are representative of the subspecies, the Yakushima macaque is much more of a dietary generalist than most primates for which there are adequate data. Am. J. Primatol. 43:305–322, 1997. © 1997 Wiley-Liss, Inc.     

Dietary differences between neighboring Cebus capucinus groups: local traditions, food availability or responses to food profitability?

The feeding patterns of three neighboring groups of Cebus capucinus were documented over a 3-year period in Santa Rosa National Park, Costa Rica. We describe the diets of the three groups and examine whether dietary differences between groups could be attributed to environmental differences in food abundances, to differences in the profitability of what was available or to learned local traditions. Diets were variable among groups; group A primarily ate fruit (81.2% of feeding time) and spent little time eating insects (16.9%), while group C was more heavily reliant on insects (44.3%) and ate less fruit (53.0%). Group B had a diet that was somewhat intermediate (69.8% fruit, 29.0% insects). By measuring the densities of all major plant foods, we were able to determine that many of the dietary differences between groups could not be attributed to simple measures of food abundance, but we could not distinguish between the food profitability hypothesis and local tradition hypothesis.     

Foraging habits of Alouatta palliata mexicana in three forest fragments

The activity patterns and diet of howler monkeys (Alouatta palliata mexicana) were studied in 3 forest fragments in south-eastern Mexico: Playa Escondida (PLA), Agaltepec Island (AGA) and Arroyo Liza (LIZ). Intersite differences offered the opportunity to investigate the foraging adaptations of howler monkeys in response to population and habitat size. In the largest fragment (PLA), the howlers' diet was based on high-quality items (fruit and young leaves). In AGA, where the density of howlers was the highest, their diet was mostly folivorous with a marked exploitation of uncommon food items such as vines, lianas, shrubs and herbs. The dietary differences in AGA were accompanied by more time spent travelling and less time spent resting. Although LIZ was the smallest fragment and had a high howler density, the small group size and the use of energy-minimizing strategies (less time spent travelling and more time spent resting) probably allowed howlers to maintain a frugivorous diet.     

The primate community of Cachoeira (Brazilian Amazonia): a model to decipher ecological partitioning among extinct species

Dental microwear analysis is conducted on a community of platyrrhine primates from South America. This analysis focuses on the primate community of Cachoeira Porteira (Para, Brazil), in which seven sympatric species occur: Alouatta seniculus, Ateles paniscus, Cebus apella, Chiropotes satanas, Pithecia Pithecia, Saguinus midas, and Saimiri sciureus. Shearing quotients are also calculated for each taxon of this primate community. Dental microwear results indicate significant differences between taxa, but are somewhat insufficient when it comes to discriminating between ecologically similar taxa. The primates of Cachoeira Porteira all incorporate a certain amount of fruit in their diet, entailing a definite amount of inter-specific competition as they must share food resources. Alouatta is the most folivorous taxon of this community, which is corroborated by dental microwear analysis. Ateles, although of a similar size to Alouatta, limits inter-specific competition by incorporating more fruit in its diet. Cebus has a very diverse omnivorous diet, which is highlighted in this study, as it compares to both fruit and leaf eating taxa. In some cases, microwear results need to be supplemented by other methods. For example, dental microwear seems insufficient to distinguish between Pithecia and Chiropotes, which eat foods with similar physical properties. However, other methods (i.e. shearing quotients and body mass) provide enough complimentary information to be able to highlight differences between the two taxa. On the other hand, dental microwear can highlight differences between primates which have similar diets, such as Saimiri and Saguinus. In this case, differences could be due to other exogenous factors.     

Morphological adaptation to diet in platyrrhine primates

Morphological features of the jaws and teeth are examined in eight species of platyrrhine monkeys that coexist in the Suriname rainforest. Z-scores calculated from geometric predictions for several features of the feeding apparatus thought to have some functional significance (e. g., tooth dimensions, jaw robusticity, leverage of primary jaw elevators) are compared to a profile of the naturalistic dietary behavior of these species (i. e., proportions of fruit mesocarp, seeds, leaves, and fauna eaten). Several features are found exclusively in those platyrrhines whose dietary preferences are the most limited. Such specializations appear to be associated with a particular protein source exploited by a species to supplement a largely frugivorous diet. Ateles paniscus, which feeds primarily on the mesocarp of ripe fruit, has an adaptive morphology that emphasizes broad incisors. Chiropotes satanas (and to a slightly lesser extent, Pithecia pithecia) is a frugivore/seed predator with large upper and lower canines and a robust mandible. The frugivore/folivore Alouatta seniculus has a relatively large total molar area and effective mandibular condyle height. In all four of these strictly vegetarian species, the leverage of the masseter muscle is greater than that of temporalis. Of the omnivorous species, Cebus apella and C. nigrivittatus exploit both fauna and seeds for protein and exhibit an array of many of the above features, such as large teeth and thick mandibles. Saimiri sciureus, not particularly known for seed predation, departs from Cebus in having less robust canines and a more gracile mandible. All three cebid omnivores have a temporalis with greater leverage than the masseter, indicating a requirement for resisting anteriorly directed forces, for example, using the jaws for vigorous foraging. The lack of any enlarged features, other than incisors, in the omnivorous Saguinus midas may be attributable to the functional constraints of small body size. Because the small size of the gape limits the size of the food parcel ingested, a requirement to enlarge other dentomandibular structures for trituration is alleviated.     

The evolution of brachiation in ateline primates, ancestral character states and history

This study examines how brachiation locomotion evolved in ateline primates using recently-developed molecular phylogenies and character reconstruction algorithms, and a newly-collected dataset including the fossils Protopithecus, Caipora, and Cebupithecia. Fossils are added to two platyrrhine molecular phylogenies to create several phylogenetic scenarios. A generalized least squares algorithm reconstructs ateline and atelin ancestral character states for 17 characters that differentiate between ateline brachiators and nonbrachiators. Histories of these characters are mapped out on these phylogenies, producing two scenarios of ateline brachiation evolution that have four commonalities: First, many characters change towards the Ateles condition on the ateline stem lineage before Alouatta splits off from the atelins, suggesting that an ateline energy-maximizing strategy began before the atelines diversified. Second, the ateline last common ancestor is always reconstructed as an agile quadruped, usually with suspensory abilities. It is never exactly like Alouatta and many characters reverse and change towards the Alouatta condition after Alouatta separates from the atelins. Third, most characters undergo homoplastic change in all ateline lineages, especially on the Ateles and Brachyteles terminal branches. Fourth, ateline character evolution probably went through a hindlimb suspension with tail-bracing phase. The atelines most likely diversified via a quick adaptive radiation, with bursts of punctuated change occurring in their postcranial skeletons, due to changing climatic conditions, which may have caused competition among the atelines and between atelines and pitheciines.     

Comparative use of color vision for frugivory by sympatric species of platyrrhines

Ateles spp. and Alouatta spp. are often sympatric, and although they are mainly frugivorous and folivorous, respectively, they consume some of the same fruit species. However, they differ in terms of color vision, which is thought to be important for fruit detection. Alouatta spp. have routine trichromatic color vision, while Ateles spp. presents the classic polymorphism of platyrrhines: heterozygous females have trichromatic color vision, and males and homozygous females have dichromatic vision. Given these perceptual differences, one might expect Alouatta spp. to consume more reddish fruits than Ateles spp., since trichromats have an advantage for detecting fruits of that hue. Furthermore, since Ateles spp. have up to six different color vision phenotypes, as do most other platyrrhines, they might be expected to include fruits with a wider variety of hues in their diet than Alouatta spp. To test these hypotheses we studied the fruit foraging behavior of sympatric Alouatta palliata and Ateles geoffroyi in Costa Rica, and modeled the detectability of fruit via the various color vision phenotypes in these primates. We found little similarity in fruit diet between these two species (Morisita = 0.086). Furthermore, despite its polymorphism, A. geoffroyi consumed more reddish fruits than A. palliata, which consumed more greenish fruits. Our modeling results suggest that most fruit species included in the diet of A. geoffroyi can be discriminated by most color vision phenotypes present in the population. These findings show that the effect of polymorphism in platyrrhines on fruit detection may not be a disadvantage for frugivory. We suggest that routine trichromacy may be advantageous for other foraging tasks, such as feeding on young leaves.     

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. J. Morphol. 275:1300–1311, 2014. © 2014 Wiley Periodicals, Inc.     

RFLP analysis of mtDNA from six platyrrhine genera: phylogenetic inferences

This study investigates the phylogenetic relationships of 10 species of platyrrhine primates using RFLP analysis of mtDNA. Three restriction enzymes were used to determine the restriction site haplotypes for a total of 276 individuals. Phylogenetic analysis using maximum parsimony was employed to construct phylogenetic trees. We found close phylogenetic relationships between Alouatta, Lagothrix and Ateles. We also found a close relationship between Cebus and Aotus, with Saimiri clustering with the atelines. Haplotype diversity was found in four of the species studied, in Cebus albifrons, Saimiri sciureus, Lagothrix lagotricha and Ateles fusciceps. These data provide additional information concerning the phylogenetic relationships between these platyrrhine genera and species.     

Seasonal Energetic Stress in a Tropical Forest Primate: Proximate Causes and Evolutionary Implications

Animals facing seasonal variation in food availability experience selective pressures that favor behavioral adjustments such as migration, changes in activity, or shifts in diet. Eclectic omnivores such as many primates can process low-quality fallback food when preferred food is unavailable. Such dietary flexibility, however, may be insufficient to eliminate constraints on reproduction even for species that live in relatively permissive environments, such as moist tropical forests. Focusing on a forest-dwelling primate with a flexible diet (Cercopithecus mitis) we investigated whether females experience seasonal energetic stress and how it may relate to reproductive seasonality. We used fecal glucocorticoids (fGCs) as an indicator of energetic stress, controlling for the potentially confounding effects of social interactions and reproductive state. We modeled within-female fGC variation with General Linear Mixed Models, evaluating changes in feeding behavior and food availability as main effects. Regardless of reproductive state, fGCs increased when females shifted their diet towards fallback foods (mature leaves and other non-preferred items) and when they spent more time feeding, while fGCs decreased with feeding time on preferred items (insects, fruits, young leaves) and with the availability of young leaves. Changes in fruit availability had no general effects on fGCs, likely because fruits were sought out regardless of availability. As predicted, females in the energetically demanding stages of late pregnancy and early lactation showed greater increases in fGCs between periods of low versus high availability of fruits and young leaves than females in other reproductive states. Potential social stressors had no measurable effects on fGCs. Preliminary evidence suggests that seasonal energetic stress may affect the timing of infant independence from mothers and contribute to unusually long inter-birth intervals compared to closely related species of similar body size. Our findings highlight how the study of stress responses can provide insights into the proximate control of reproductive strategies.     

Comparative and functional myology of the prehensile tail in new world monkeys

The caudal myology of prehensile-tailed monkeys (Cebus apella, Alouatta palliata, Alouatta seniculus, Lagothrix lagotricha, and Ateles paniscus) and nonprehensile-tailed primates (Eulemur fulvus, Aotus trivirgatus, Callithrix jacchus, Pithecia pithecia, Saimiri sciureus, Macaca fascicularis, and Cercopithecus aethiops) was examined and compared in order to identify muscular differences that correlate with osteological features diagnostic of tail prehensility. In addition, electrophysiological stimulation was carried out on different segments of the intertransversarii caudae muscle of an adult spider monkey (Ateles geoffroyi) to assess their action on the prehensile tail. Several important muscular differences characterize the prehensile tail of New World monkeys compared to the nonprehensile tail of other primates. In atelines and Cebus, the mass of extensor caudae lateralis and flexor caudae longus muscles is more uniform along the tail, and their long tendons cross a small number of vertebrae before insertion. Also, prehensile-tailed monkeys, especially atelines, are characterized by well-developed flexor and intertransversarii caudae muscles compared to nonprehensile-tailed primates. Finally, Ateles possesses a bulkier abductor caudae medialis and a more cranial origin for the first segment of intertransversarii caudae than do other prehensile-tailed platyrrhines. These myological differences between nonprehensile-tailed and prehensile-tailed primates, and among prehensile-tailed monkeys, agree with published osteological and behavioral data. Caudal myological similarities and differences found in Cebus and atelines, combined with tail-use data from the literature, support the hypothesis that prehensile tails evolved in parallel in Cebus and atelines. © 1995 Wiley-Liss, Inc.     

Functional patterns of molar occlusion in platyrrhine primates

Mechanico-functional features of molar form were studied in Callithrix, Alouatta, Pithecia and Cebus. Molars of Callithrix and Alouatta are adapted to loading foods under relatively high occlusal pressure; those of Pithecia and Cebus, under relatively low occlusal pressure. General functional considerations suggest that these taxa are adapted to insectivorous, folivorous, frugivorous and omnivorous diets, respectively. The physical properties of foods, principally mechanical strength and deformability, determine the selective pressures involved in the evolutionary adaptation of molar form. A dietary classification based upon percentages of foods eaten does not always reflect morphological adaptations. Homologous parts of teeth and homologous parts of the masticatory cycle do not always subserve equivalent functions. The relevance of functional occlusal analysis for deciphering phylogeny and explaining evolutionary grades is stressed.     

Diets of some French guianan primates: Food composition and food choices

We studied food choices of black spider monkeys (Ateles paniscus) and red howlers (Alouatta seniculus) in an undisturbed tropical forest of French Guiana for 6 months in the rainy season. We made additional observations on tufted capuchins (Cebus apella) and examined the differences and similarities in feeding behavior with respect to the plant specific composition of the habitat and the biochemical characteristics of their food. Capuchins and spider monkeys mainly fed on ripe fruit pulp, to which they added invertebrates (capuchins) or young leaves (spider monkeys); their plant diet was more varied than that of howlers, which contained approximately equal proportions of ripe fruits and young leaves of many species, with large monthly variations in these food categories. BothAteles andAlouatta tended to feed preferentially on abundant plant species, with a large overlap in their fruit choices. The former species included a high proportion of soluble sugars and a low proportion of protein in its diet compared, to howlers. Leaves of several species selected byAlouatta reacted positively when screened for alkaloids and phenolic compounds. Besides specific metabolic requirements and adaptations to deriving nutrients from distinct food types, we hypothesize that the ability to taste sugars, which varies among primates, affects the range of foods appearing to be palatable and, consequently, contributes to the differentiation of feeding niches.     

Differential habitat use byCebus apella andSaimiri sciureus in central surinam

Cebus apella, the black-capped or tufted capuchin, andSaimiri sciureus, the squirrel monkey, are frequently found in mixed species feeding and foraging groups throughout tropical South America and have been reported previously to show marked differences in techniques of insect foraging. Individuals of these taxa observed in central Surinam show numerous additional species-specific differences in other aspects of habitat utilization including gross dietary preferences, utilization of forest strata, locomotor behavior and arboreal support preferences.Cebus apella is more frugivorous, frequents the middle and lower levels of the main canopy, is predominantly quadrupedal and moves on medium-sized arboreal supports. By contrast, the smallerSaimiri sciureus is more insectivorous, frequents the lower strata of the forest, is more saltatory and moves on the smallest arboreal supports. Many of these differences in habitat use are interrelated and accord with patterns of habitat use seen in other primate taxa. The differences betweenCebus apella andSaimiri sciureus also accord with the types of behavioral differences frequently associated with differences in body size.     

Diet, phylogeny, and basal metabolic rate in phyllostomid bats

Aside from the pervasive effects of body mass, much controversy exists as to what factors account for interspecific variation in basal metabolic rates (BMR) of mammals; however, both diet and phylogeny have been strongly implicated. We examined variation in BMR within the New World bat family Phyllostomidae, which shows the largest diversity of food habits among mammalian families, including frugivorous, nectarivorous, insectivorous, carnivorous and blood-eating species. For 27 species, diet was taken from the literature and BMR was either measured on animals captured in Brazil or extracted from the literature. Conventional (nonphylogenetic) analysis of covariance (ANCOVA), with body mass as the covariate, was first used to test the effects of diet on BMR. In this analysis, which assumes that all species evolved simultaneously from a single ancestor (i.e., a "star" phylogeny), diet exerted a strong effect on mass-independent BMR: nectarivorous bats showed higher mass-independent BMR than other bats feeding on fruits, insects or blood. In phylogenetic ANCOVAs via Monte Carlo computer simulation, which assume that species are part of a branching hierarchical phylogeny, no statistically significant effect of diet on BMR was observed. Hence, results of the nonphylogenetic analysis were misleading because the critical values for testing the effect of diet were underestimated. However, in this sample of bats, diet is perfectly confounded with phylogeny, because the four dietary categories represent four separate subclades, which greatly reduces statistical power to detect a diet (= subclade) effect. But even if diet did appear to exert an influence on BMR in this sample of bats, it would not be logically possible to separate this effect from the possibility that the dietary categories differ for some other reason (i.e., another synapomorphy of one or more of the subclades). Examples such as this highlight the importance of considering phylogenetic relationships when designing new comparative studies, as well as when analyzing existing data sets. We also discuss some possible reasons why BMR may not coadapt with diet.     

Diet and activity in black howler monkeys (<Emphasis Type="Italic">Alouatta pigra</Emphasis>) in southern Belize: does degree of frugivory influence activity level?

This study reports on the diet and activity budgets of Central American black howling monkeys (Alouatta pigra) at Monkey River, Belize. This is a previously unstudied population, close to the southern boundary of the species range, and it provides comparative data on A. pigra from a new study site. Both diet and activity are within the ranges reported for other A.pigra sites and for mantled howlers (A. palliata). No age-sex differences could be discerned in either diet or activity, though monthly variation was apparent. The monkeys switch from consuming leaves 86% of the time in January to March to consuming 67% fruit in April to July. This difference was statistically significant, and provided the opportunity to compare activity levels of the monkeys over two dietary periods, one characterized primarily by folivory, the other by frugivory. Howlers are often seen as a relatively inactive species, something that is associated with a low quality, folivorous diet. However, A. pigra have been described as being as frugivorous as possible and as folivorous as necessary. Yet, despite the opportunistic consumption of large quantities of high-energy foods, A. pigra has been observed as conforming to the howler lifestyle, resting as much as 80% of the day. The data in this paper support both of these reports. Black howlers at Monkey River Belize are typically inactive, maintaining high levels of inactivity even during months characterized by frugivory, suggesting that diet is more flexible and varied than is behavior and calling into question the assumption that howler inactivity is due to the digestion of large quantities of leaves.