Effects of Food Type and Number of Feeding Sites in a Tree on Aggression During Feeding in Wild <Emphasis Type="Italic">Macaca fuscata</Emphasis>

It is important to understand the effects of ecological factors on aggression during feeding in order to link habitat characteristics to competitive regime and social relationships. Multiple habitat characteristics are likely to affect aggression, but few researchers have examined the effect of multiple factors on intragroup competition simultaneously. I examined the effect of 8 factors on aggression during feeding in wild Japanese macaques living in a coniferous forest in Yakushima: density of the tree species, feeding time, number of feeding sites within a feeding tree, number of cofeeding animals, intratree macaque density, food type, and rank and age of the focal individual. When macaques cofed with other individuals, food type, the number of feeding sites, and their interactions significantly influenced aggression. Aggression increased when macaques ate fruits/seeds versus other foods and as the number of feeding sites decreased. Primate socioecological models highlight the importance of clumped distribution of food patches as a correlate of intragroup contest. However, my study indicates that primatologists need to pay attention to the factors related to the current feeding tree—food type and feeding tree size with respect to monopolizability—in addition to the distribution of food in the entire home range.     

Social feeding decisions in horses (Equus caballus)

Like many other herbivores, in a natural environment equids feed on rather evenly distributed resources. However, the vegetation in their vast habitats constantly changes. If food is plentiful only little competition occurs over food, and in non-competitive situations domestic horses tend to return to the same feeding site until it is overgrazed. In contrast, they compete over limited food for which the social status of the individuals appears to be important. Especially in ruminants several studies have proved an influence of social organisations, rank, sex and the depletion of feeding sites on the feeding behaviour of individuals. However, it is not yet understood whether and how social aspects affect horses' feeding decisions. Curiosity about the influence of social rank on the horses' feeding decisions between two, equally with high-quality surplus food-filled buckets placed in different social feeding conditions, led us to create the test below. The observer horses were alternately tested with a dominant and a subordinate demonstrator placed in one of the three different positions. We conclude that domestic horses use social cognition and strategic decision making in order to decide where to feed in a social feeding situation. When possible they tend to return to the same, continuously supplied feeding site and switch to an "avoidance tendency" in the presence of dominant horses or when another horse is already feeding there. Thus, the social rank and the position of conspecifics affect the feeding strategy of horses.     

On the feeding and social behaviour of the laying hen

This paper describes a study of the influences of early husbandry conditions, social attraction and social rank on various aspects of the feeding behaviour of laying hens.Birds were raised in flocks of 10, 60 or 500. Groups of 3 birds, selected from flocks of the same size, were then housed in pens. Some groups consisted of hens raised in the same flock, and some of birds raised in different flocks. The feeding and agonistic behaviour of each group at each of 5 types of feeder was observed and compared with the behaviour shown when the birds had free access to a 1-m long food trough. Each of the 5 feeders offered the same area of feeding space, but differed in its partitioning and spatial distribution. One feeder had a single unpartitioned feeding space. The other 4 feeders had 3 partioned feeding spaces which were adjacent, or separated by distances of 10, 20 or 40 cm, respectively.For the 5 feeders, total feeding times and lengths of feeding bouts were greatest, and the number of feeding bouts least, when the feeding space was unpartitioned. Synchrony of feeding behaviour was low when the feeding space was unpartitioned or the partitioned spaces adjacent, but was comparable to that at the 1-m long food trough when the distance between partitioned feeding spaces was 10 cm or greater. When feeding space was partitioned, the likelihood that 2 birds would eat together at the same site increased with the distance between feeding space. Dominant birds always exhibited the longest feeding bouts and greatest total feeding times, but were less likely to feed in the same space as another bird, and exhibited less synchrony of feeding behaviour than subordinates. The size of the flock in which the birds were raised, and whether or not the birds in a group had been raised together or apart, had no clear effect on behaviour.These results indicate that, within the limits of this experiment, early husbandry conditions do not influence behaviour shown during feeding in later life, and that social attraction has a greater influence on the feeding behaviour of hens than is generally assumed. In view of this latter finding, it is postulated that in attempting to determine the requirements of laying hens for feeding space, attention must be paid to social attraction as well as to competition at the feeder.     

Decisions on time allocation to different food patches by Japanese monkeys (Macaca fuscata)

From the standpoint of the possible development of an optimal foraging strategy, the time allocation to different food patches and the decision as to when to leave any given patch were examined in wild Japanese monkeys (Macaca fuscata). The investigations yielded the following results: (1) There was no tendency for feeding bouts to be of longer duration in higher-quality patches, i.e. the monkeys did not employ an energy-maximizing strategy during their feeding bouts. (2) The speed of feeding barely decreased with passage of time during a bout. In most cases, therefore, a decrease in feeding speed was not considered relevant to the decision to leave a patch. These findings are evaluated in terms of social factors, and it appears that following a monkey's own family group or the troop usually contributes to the timing of leaving a food patch.     

A lifetime perspective on foraging and mortality

Food intake carries many potential risks which may impair an animal's reproductive success not only in the current breeding cycle, but also for the rest of its lifetime. We examine the lifetime trade-off between the costs and benefits of food intake by presenting a simple animal foraging model, where each unit of food eaten carries with it a risk of mortality. We show that the optimal food intake rate over an animal's lifetime, for both semelparous and iteroparous animals, is not maximal. Instead, animals are required to strike a balance between the immediate reproductive benefits of gathering food and the future reproductive costs incurred by the food's mortality risk. This balance depends upon the lifespan of the animal as well as the nature of the risk. Different mortality risks are compared and it is shown that a mortality risk per unit time spent foraging is not, in general, equivalent to a mortality risk per unit of food consumed. The results suggest that a mortality risk per unit of food consumed, such as that presented by the presence of a toxin or of a parasite in the diet, has important consequences for feeding behaviour and is a possible factor involved in food intake regulation.     

Behavioural consequences of hunting by expectation: A simulation study of foraging tactics

Foraging behaviour has been simulated using a model that predicts the path taken by an animal foraging for particulate food, the path being defined by the animal's remembrance of its previous foraging success. This is represented in the model by a two-dimensional vector with its modulus encoding an exponentially smoothed average of the animal's feeding rate and its orientation encoding the average direction of travel. As food is ingested the amount ingested and its position are used to update the remembrance, and the animal turns in the adjusted direction travelling at a speed inversely proportional to the average feeding rate. Foraging paths simulated in a patchy environment are shown to have the following properties: (i) They tend to avoid crossing the boundaries of patches from the inside. (ii) They tend to avoid intersecting themselves. (iii) When they do intersect themselves they usually do so more or less at right angles. (iv) They ascend gradients of density of food.     

Use of sleeping trees by ursine colobus monkeys (Colobus vellerosus) demonstrates the importance of nearby food

Examination of the characteristics and locations of sleeping sites helps to document the social and ecological pressures acting on animals. We investigated sleeping tree choice for four groups of Colobus vellerosus, an arboreal folivore, on 298 nights at the Boabeng-Fiema Monkey Sanctuary, Ghana using five non-mutually exclusive hypotheses: predation avoidance, access to food, range and resource defense, thermoregulation, and a null hypothesis of random selection. C. vellerosus utilized 31 tree species as sleeping sites and the species used differed per group depending on their availability. Groups used multiple sleeping sites and minimized their travel costs by selecting trees near feeding areas. The percentage that a food species was fed upon annually was correlated with the use of that species as a sleeping tree. Ninety percent of the sleeping trees were in a phenophase with colobus food items. Entire groups slept in non-food trees on only one night. These data strongly support the access to food hypothesis. Range and resource defense was also important to sleeping site choice. Groups slept in exclusively used areas of their home range more often than expected, but when other groups were spotted on the edge of the core area, focal groups approached the intruders, behaved aggressively, and slept close to them, seemingly to prevent an incursion into their core range. However, by sleeping high in the canopy, in large, emergent trees with dense foliage, positioning themselves away from the main trunk on medium-sized branches, and by showing low rates of site reuse, C. vellerosus also appeared to be avoiding predation in their sleeping site choices. Groups left their sleep sites later after cooler nights but did not show behavioral thermoregulation, such as huddling. This study suggests that access to food, range and resource defense, and predation avoidance were more important considerations in sleeping site selection than thermoregulation for ursine colobus.     

Feeding rates of a mammalian browser confirm the predictions of a ‘foodscape’ model of its habitat

Adequate nutrition is a fundamental requirement for the maintenance and growth of populations, but complex interactions between nutrients and plant toxins make it difficult to link variation in plant quality to the ecology of wild herbivores. We asked whether a ‘foodscape’ model of habitat that uses near-infrared spectroscopy to describe the palatability of individual trees in the landscape, predicted the foraging decisions of a mammalian browser, the koala (Phascolarctos cinereus). Specifically, we considered four behavioural decision points at which nutritional quality may influence an animal’s decision. These were: which tree to enter, whether to feed from that tree, when to stop eating, and how long to remain in that tree. There were trends for koalas to feed in eucalypt trees that were more palatable than unvisited neighbouring conspecific trees, and than trees that they visited but did not eat. Koalas ate longer meals in more palatable trees, and stayed longer and spent more time feeding per visit to these trees. Using more traditional chemical analyses, we identified that an interaction between the concentrations of formylated phloroglucinol compounds (a group of plant secondary metabolites) and available N (an integrated measure of tannins, digestibility and N) influenced feeding. The study shows that foodscape models that combine spatial information with integrated measures of food quality are a powerful tool to predict the feeding behaviour of herbivores in a landscape.     

Measuring social tolerance: An experimental approach in two lemurid primates

Social tolerance crucially affects the life of group‐living animals as it can influence, among other things, their competitive regimes, access to food, learning behavior, and recruitment. However, social tolerance tests were mainly conducted in semi‐free or captive populations, and we know little about the behavioral mechanisms and consequences of social tolerance under natural conditions. We therefore developed a co‐feeding experiment to measure social tolerance in groups of wild and captive animals across two primate species. Specifically, we recorded the social tolerance level of redfronted lemurs (Eulemur rufifrons, four wild, one captive group) and ringtailed lemurs (Lemur catta, three wild, three captive groups) by presenting a clumped food resource in an experimental arena, and recorded patterns of resource use during the experiment. Because redfronted lemurs exhibit lower levels of decided conflicts than ringtailed lemurs, we predicted that they would be socially more tolerant. The probability for an individual to feed in the arena was higher in redfronted lemurs than in ringtailed lemurs. In addition, in both species, the probability for an individual to feed in the arena was higher in the captive populations than in their wild counterparts, suggesting that proximate factors, such as a relaxation of feeding competition in captivity, may adapt species‐specific levels of social tolerance to local levels of food availability. Hence, the number of individuals co‐feeding on a valuable food resource appears to be a useful proxy of social tolerance that could be measured with this experimental setup in other wild and captive species as well.     

State-dependent ideal free distributions

Summary The standard ideal free distribution (IFD) states how animals should distribute themselves at a stable competitive equilibrium. The equilibrium is stable because no animal can increase its fitness by changing its location. In applying the IFD to choice between patches of food, fitness has been identified with the net rate of energetic gain. In this paper we assess fitness in terms of survival during a non-reproductive period, where the animal may die as a result of starvation or predation. We find the IFD when there is a large population that can distribute itself between two patches of food. The IFD in this case is state-dependent, so that an animal's choice of patch depends on its energy reserves. Animals switch between patches as their reserves change and so the resulting IFD is a dynamic equilibrium. We look at two cases. In one there is no predation and the patches differ in their variability. In the other, patches differ in their predation risk. In contrast to previous IFDs, it is not necessarily true that anything is equalized over the two patches.     

The abundance,distribution and structural characteristics of tree‐holes in Nothofagus forest,New Zealand

Tree‐holes provide an important microhabitat that is used for feeding, roosting and breeding by numerous species around the world. Yet despite their ecological importance for many of New Zealand's endangered species, few studies have investigated the abundance or distribution of tree‐holes in native forests. We used complementary ground and climbed tree surveys to determine the abundance, distribution and characteristics of tree‐holes in undisturbed Nothofagus forest in the Lewis Pass, New Zealand. We found that hole‐bearing trees were surprisingly abundant compared with many other studies, including Australian Eucalyptus species and American beech. In fact, we estimated as many as 3906 tree‐holes per hectare, of which 963 holes per hectare were potentially large enough to provide roost sites for hole‐nesting bats in New Zealand, while only eight holes per hectare were potentially suitable for specialist hole‐nesting birds. This was of great interest as primary cavity‐excavating animals are absent from New Zealand forests, compared with North America and Australia. Moreover, tree‐hole formation in New Zealand is likely to be dominated by abiotic processes, such as branch breakage from windstorms and snow damage. As has been found in many other studies, tree‐holes were not uniformly distributed throughout the forest. Tree‐holes were significantly more abundant on the least abundant tree species, Nothofagus fusca, than on either N. menziesii or N. solandri. In addition to tree species, tree size was also an important factor influencing the structural characteristics of tree‐holes and their abundance in this forest. Moreover, these trends were not fully evident without climbed tree surveys. Our results revealed that ground‐based surveys consistently underestimated the number of tree‐holes present on Nothofagus trees, and illustrate the importance of using climbed inspections where possible in tree‐hole surveys. We compare our results with other studies overseas and discuss how these are linked to the biotic and abiotic processes involved in tree‐hole formation. We consider the potential implications of our findings for New Zealand's hole‐dwelling fauna and how stand dynamics and past and future forest management practices will influence the structural characteristics of tree‐holes and their abundance in remnant forest throughout New Zealand.     

Predicting the frequency of food-related agonism in white-faced capuchin monkeys (Cebus capucinus), using a novel focal-tree method

Food abundance and distribution have played a central role in the conceptual theory of primate socioecology [Janson, Behaviour 105:53-76, 1988; Isbell, Behavioral Ecology 2:143-155, 1991; Sterck et al., Behavioral Ecology and Sociobiology 41:291-309, 1997; van Schaik, In: Standen V, Foley RA, editors, Comparative Socioecology. Oxford: Blackwell. p 195-218, 1989]. This theory predicts that agonistic ("contest") competition should occur when food is distributed in discrete, defensible patches; in contrast, when food sources are distributed uniformly or randomly, non-agonistic ("scramble") competition is expected. Primatologists usually measure resource density and patchiness from a botanical perspective, ignoring the biology of the animal being studied. Such an approach may be irrelevant in terms of how animals view the dispersion of resources. Using a novel focal-tree method that measures resource availability on a scale that is both spatially and temporally relevant to the animal under investigation, we take a cost-benefit approach to predict the frequency of food-related agonism in white-faced capuchin monkeys (Cebus capucinus) from 11 ecological and social variables. We retained four variables in the regression model: two representing the opportunity for aggression (i.e., feeding bout length and the number of feeding adult females), and two representing opportunity costs (i.e., fruit abundance and the number of potential feeding sites in the focal tree). The results of this study indicate that the amount of food-related aggression in white-faced capuchins can be predicted by variables representing the costs and benefits of contesting a food resource.     

Grazing behaviour of sheep in a situation of conflict between feeding and social motivations

We investigated how food preferences and social bonds interact to determine the choice of grazing location in sheep. Ewes of INRA 401 breed were grazed in plots in which taller areas, i.e. preferred feeding sites, were left to grow at 15 or 50 m from a socially attractive site, i.e. familiar ewes placed in a public pen at one end of the plot. Eight experimental ewes were tested either alone or in groups with one, three or six accompanying animals chosen amongst 20 other familiar ewes. We used a Latin square design, in which the eight treatments (two distancesxfour group sizes) were balanced in 8 measurement days. We recorded, in 20-min tests, the behaviour of the experimental ewes by focal sampling, and the location of each animal in the groups by scan sampling. Foraging location, dietary choices and vigilance behaviour of ewes were affected by both the distance between the group of public peers and the preferred feeding site, and the size of their own group. Our results suggest that a sheep will move whether alone or with a few peers to a preferred feeding site located close to the core of its social group. In a small sub-group, its frequency of vigilance behaviour increases, probably to maintain social contact with the rest of the group. Conversely, a sheep will not leave its group to reach a preferred feeding site located further away unless it is followed by several other peers.     

The approximately ideal, more or less free distribution

We present the minimum set of requirements necessary and sufficient to represent the foraging behaviour of an animal, and its utilisation of food, in order to explore the emergent properties of behaviour that allow animals to reduce their hunger. We present an individual-based model of foraging that provides a simple quantification of the requirements, which is sufficiently simple to yield some analytical results. Complex interactions beyond the scope of analysis have been explored through simulating animals foraging in regenerating patchy environments. In most cases the populations pass into equilibrium distributions which appear to be stable. The equilibria always approximate closely to the ideal free distribution, although typically with a small degree of undermatching. (Undermatching is the term applied to the departure from the ideal free distribution caused by a smaller proportion of the population than expected occupying areas with a higher than average regeneration rate). The model therefore implies that the distribution, hitherto accounted for in terms of ESSs may, in fact, be simply an effect of the animal's utilization of the food it collects to reduce its hunger. The model defines a specific feeling rate, v, the rate at which an animal can feed on a unit of food. This is a function of three parameters, v1, the specific feeding rate when alone, v(infinity), the rate, possibly zero, at which it can feed in the presence of an indefinitely large number of conspecifics, and n1/2, the number of conspecifics that cause v to take the value (v1+v(infinity)/2. Exploitation competition in the absence of interference is represented by setting v1 = v(infinity). Differences in competitive ability in exploitation have been represented by simulating animals with a range of values of v1, those with the larger values, feeding more rapidly, being the more effective competitors, and those with the lower values being the less effective. Interference competition is represented by setting v1 > v(infinity) and social facilitation by v1 < v(infinity). Individual differences in the strength of interaction are represented by different values of n(1/2). In competition, the animals with the larger values of n(1/2) are the more effective competitors: in facilitation, they are the less effective facilitators. The addition of physiological and behavioural detail makes very little alteration to the emergent equilibria, always close to the ideal free distribution, almost always showing undermatching.     

Time and energy constraints in demanding phases of the annual cycle: an example of time limitation in refuelling migratory swans

The daily metabolizable energy intake of an animal is potentially limited by either the available feeding time or by its capacity to process energy. Animals are generally considered not to be time-limited but rather to be energy-processing-limited. This is concluded from the common observation that an animal's feeding time per day increases with a decrease in food density. We argue that such changes in feeding time are in theory also expected when no constraints are operating. Thus, a study of the constraints on energy intakes of free-living animals should be performed during demanding phases of the year. As an example, we collected data on time and energy budgets of Bewick's swan ( Cygnus columbianus bewickii ) refuelling during migration on fennel pondweed ( Potamogeton pectinatus ) tubers in two years differing two-fold in tuber biomass density. As predicted by time limitation, the feeding time (defined as the time with the head submerged) did not change in response to a change in food biomass density, both within and between years (averaging 12.2 h d⁻¹). Contrary to energy-processing limitation, and again in line with time limitation, the daily metabolizable energy intake varied, being greater in the year with high than in the year with low food densities. We conclude that more studies are needed of animals operating under demanding conditions before it can be assessed whether free-living animals are generally energy-processing- or time-limited.     

Costs and benefits of within-group spatial position: a feeding competition model

An animal's within-group spatial position has several important fitness consequences. Risk of predation, time spent engaging in antipredatory behavior and feeding competition can all vary with respect to spatial position. Previous research has found evidence that feeding rates are higher at the group edge in many species, but these studies have not represented the entire breadth of dietary diversity and ecological situations faced by many animals. In particular the presence of concentrated, defendable food patches can lead to increased feeding rates by dominants in the center of the group that are able to monopolize or defend these areas. To fully understand the tradeoffs of within-group spatial position in relation to a variety of factors, it is important to be able to predict where individuals should preferably position themselves in relation to feeding rates and food competition. A qualitative model is presented here to predict how food depletion time, abundance of food patches within a group, and the presence of prior knowledge of feeding sites affect the payoffs of different within-group spatial positions for dominant and subordinate animals. In general, when feeding on small abundant food items, individuals at the front edge of the group should have higher foraging success. When feeding on slowly depleted, rare food items, dominants will often have the highest feeding rates in the center of the group. Between these two extreme points of a continuum, an individual's optimal spatial position is predicted to be influenced by an additional combination of factors, such as group size, group spread, satiation rates, and the presence of producer-scrounger tactics.     

Social models fail to induce diet and feeding site avoidance in naïve yearling steers

Social learning can be of critical importance to cattle grazing rangeland environments with high variability of food resources across space and time. Experienced individuals can greatly facilitate foraging decisions (what to eat and where to eat) of na?ve peers in such settings. We conducted an experiment with cattle to investigate strength and persistence of socially induced food and feeding site avoidance behaviours. Sixteen na?ve yearling steers were paired with 16 social models that had either not been trained (control) or been trained with an emetic (LiCl), electrical shock or both to avoid: (a) an unsafe high-quality food (LiCl); (b) an unsafe high-quality feeding site (shock); or (c) both the unsafe high-quality food and the unsafe high-quality feeding site (LiCl + shock). Ten-minute trials were conducted in an experimental arena containing three artificial feeding sites each consisting of groups of bowls with either high- (HQ) or moderate-quality (MQ) foods (HQ = barley and oat grain; MQ = Bermuda grass hay). Unsafe high-quality (UHQ, surrounded by traffic cones) and safe moderate-quality (SMQ) feeding sites consisted of nine rubber bowls containing either HQ or MQ foods. The safe high-quality (SHQ) feeding site consisted of two groups of eight bowls containing HQ food, which surrounded the UHQ and SMQ feeding sites. Social models did not induce diet and feeding site avoidance behaviours in na?ve steers; they exerted small and transient changes in the feeding behaviour of their na?ve counterparts. Consequences to the individual outweighed social influences; when na?ve animals experienced the same punishment contingencies as their social models, their behavioural patterns closely resembled those of their social model. Conditioned food and location aversions via LiCl were apparently influenced by prior exposure to target foods and the experimental arena. Conversely, conditioned feeding site avoidance via shock was apparently not influenced by prior exposure to target foods or the experimental arena.     

Social facilitation does not reduce neophobia in Chestnut-sided Warblers (Parulinae:Dendroica pensylvanica)

Chestnut-sided Warblers (Dendroica pensylvanica) show aversion to feeding at a familiar food source if placed next to a novel object. I have suggested that neophobia is an adaptation of specialized animals to reduce wasteful exploration. In this experiment I tested whether social facilitation mitigates the fear and decreases the latency to forage at a novel foraging site. As far as I know, the effect of social factors on neophobia has not been previously investigated. Because Chestnut-sided Warblers are intraspecifically solitary during the non-breeding season, I examined their response to novel foraging sites in the presence of a Bay-breasted Warbler (D. castanea) “trainer”. Chestnut-sided Warblers commonly join flocks with this more generalized and less neophobic congener. Chestnut-sided Warblers showed no tendency to feed in the presence of a Bay-breasted Warbler that is freely feeding from the unfamiliar microhabitat. Specialized animals may be more resistant to social facilitation and observational learning than more generalized species.     

An analysis of dyadic interactions of male Japanese monkeys (Macaca fuscata fuscata) in a cage-room observation

This is a laboratory observation (Home-cage and observation room) for individual and social behavior of four male Japanese monkeys. The monkeys were placed in a dyadic and trio situation with the purpose to investigate the relationships between aggressive and affinitive factors which regulated their social behaviour. Total amounts of attacks and receiving groom showed a consistent order for four monkeys through the pairwise conditions, which was presumed to indicate a dominance order. Grooming and mounting behavior, however, were influenced by the particular dyadic interactions, probably social attraction, rather than dominance order itself. The presence of the third animal facilitated such a social preference between animals positioned closely in the rank order as well as elicited aggression from the dominant animals.     

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.