Transition from a random to an ideal free to an ideal despotic distribution: the effect of individual difference in growth

Individual differences in growth can lead to a monopolistic form of food competition. We studied the long-term transition in the mode of competition and the distribution of individuals between food patches of the cloned salmonid fish, Oncorhynchus masou ishikawae, in the laboratory. This transition was accompanied by growth depensation, i.e., the increase over time in the variance of size between individuals resulting from the differences in individual growth rates. The 120-cm experimental tanks were divided into two compartments (patches) between which an opaque partition was placed. Fish were able to move freely between the patches and therefore were able to assess the patch quality using long-term memory, but they were not able to see the food input in the other patch directly. The distribution between the two food patches, the amount of food gained, and the growth and the agonistic behavior of four groups of six individuals were observed over 4 weeks. We found that (1) within-group variation in body weight increased with time; (2) on average, the better patch was used by more individuals than predicted by a random distribution but fewer individuals than predicted by an ideal free distribution, and (3) the distribution and pattern of resource use by the fish changed over the 4-week experimental period from a random distribution to an ideal free distribution and finally to an ideal despotic distribution. We suggest that growth depensation causes the long-term change in the spatial distribution and pattern of resource use by competitors. Received: December 19, 2000 / Accepted: March 19, 2001     

Test of the ecological-constraints model on ursine colobus monkeys (Colobus vellerosus) in Ghana

For group-living mammals, the ecological-constraints model predicts that within-group feeding competition will increase as group size increases, necessitating more daily travel to find food and thereby constraining group size. It provides a useful tool for detecting scramble competition any time it is difficult to determine whether or not food is limiting. We tested the ecological-constraints model on highly folivorous ursine colobus monkeys (Colobus vellerosus) at the Boabeng-Fiema Monkey Sanctuary in Ghana. Three differently sized groups were followed for 13 months and two others were followed for 6 months each in 2004-2005 using focal-animal sampling and ranging scans; ecological plots and phenology surveys were used to determine home-range quality and food availability. There was relatively little difference in home-range quality, monthly food availability, diet, adult female ingestion rates, and rate of travel within food patches between the groups. However, home-range size, day-range length, and percent of time spent feeding all increased with group size. We performed a single large test of the ecological-constraints model by combining several separate Spearman correlations, each testing different predictions under the model, using Fisher's log-likelihood method. It showed that the ecological-constraints model was supported in this study; scramble competition in this population is manifesting in increased ranging and time spent feeding. How costly this increased energy expenditure is for individuals in larger groups remains to be determined.     

The influence of social rank in the angelfish,Pterophyllum scalare,on locomotor and feeding activities in a novel environment

To investigate differences in behaviour associated with social rank and environmental conditions, a comparison was made between swimming and feeding activities of dominant and subordinate angelfish, housed in groups of six, with those of angelfish housed singly in identical laboratory aquaria. Subordinate individuals were less active, less ready to feed and consumed less food items than dominants, but their feeding and activity levels were still greater than those displayed by isolated fish used as controls. When fish from each of the above categories were transferred to a novel, identical tank to be tested individually for a 6-day period, little change was observed in previously isolated fish. In contrast, the previous social experience had a marked influence on the behaviour of the other individuals, the effects being related to the social status. The strongest initial response to the new environment was shown by subordinate individuals, with a significant increase in swimming and a significant decrease in feeding compared to the group situation. Subordinates were significantly more active than dominants, who in turn moved more than previously isolated fish. Dominants and subordinates were now similarly reluctant to feed, and their food consumption was less than that of previously isolated fish. With time in the novel environment a significant reduction of swimming activity and a recovery of feeding measures were detected, but levels were still depressed in relation to the group condition, and lower in subordinates than in dominants, indicating the long-lasting effects of the previous social interactions. The results have clear implications for laboratory studies using groups of fish in which social hierarchies may be established prior to individuals being tested singly in a novel environment.     

Competition for Resources and Its Behavioral Consequences Among Female Primates

Via the current model on the evolutionary ecology of female social relationships, Sterck et al. (1997) argue that ecological conditions determine how competition over food resources affects female fitness. The relative importance of different modes of competition then affects female social relationships and dispersal patterns. I outline the model and review relevant data. There are 3 modes of feeding competition: within-group scramble (WGS), within-group contest (WGC), and between-group contest (BGC), which occur in various combinations in different populations of nonhuman primates. Ecological measures support predictions that limiting resources lead to WGS and clumped resources induce WGC. The ecological basis of BGC remains elusive, but it is probably linked to resource abundance. Tests of the proxies of feeding competition support the idea that short-term search substrates and increasing group size lead to WGS, while high-quality patches of intermediate size relative to group size lead to WGC. However, when tested across populations, independent measures of aggression rates do not always match the actual or presumed competitive regimes. This mismatch might be explained by confounding factors and the predominately indirect measures of feeding competition. Predicted relationships between feeding competition and female social relationships/dispersal are only partly supported. This might be attributed to the fact that few studies have taken ultimate approaches using mechanistic correlates of fitness (net energy gain) or lifetime reproductive success to measure consequences of feeding competition. But to resolve existing inconsistencies, additional factors need to be taken into account as well, for example, male sexual strategies may affect female feeding competition; constraints on group size may enforce female dispersal; and demography may alter rates of alliances. More explicitly, ultimate approaches are needed to test the consistency of the socioecological model.     

The Effect of Within-Flock Spatial Position on the Use of Social Foraging Tactics in Free-Living Tree Sparrows

The benefit of producer (searches for own food) or scrounger (exploits the others’ food discoveries) foraging tactic in a group of socially feeding animals may depend on where the individual searches for food within the group. Scrounging may be more advantageous in the centre of the group, having more individuals around to join, while producing may be more beneficial at the edges, where more unexplored food patches may be found. This study shows within‐flock position correlates with foraging tactic use of feeding birds in socially foraging tree sparrows, Passer montanus. Sparrows staying closer to the centre of the flock found their food patches more frequently by joining (i.e. use more frequently the scrounging tactic) than those staying toward the edges. To our knowledge this is the first field study demonstrating the relationship between spatial position and foraging tactic use. We investigated this relationship under different perceived predation hazard, and found that under elevated risk of predation, central individuals may increase their use of joining more than individuals on the periphery of the flock. Moreover, we show that extremely specialized use of searching tactics may be very infrequent in tree sparrows. As both within‐flock position and search tactic use can be altered very quickly and without leaving the flock, individuals may easily alter them in order to adjust their behaviour.     

Exploitation of food resources by the cockroach Blattella germanica in an urban habitat

The exploitation of food resources by the German cockroach, Blattella germanica (L.) (Dictyoptera: Blattellidae) was investigated experimentally in relation to distance from shelters and depletion of neighbouring food patches. In addition, the dynamics of exploitation of a patch were analysed. Observations were made after dark in a public swimming baths building and each one lasted 3 h. Food patches were placed in rows, at different distances from the shelters. The number of cockroaches in food dishes, in a 20 cm diameter circle round each food dish and in a 60 cm diameter circle round this first circle were recorded.Food items nearest the shelters were exploited first. Exploitation of row 2 and of row 3 food items started later, after row 1 food patches had been depleted. Under these conditions, the moment a food patch was exploited was related to its distance from shelter. Exploitation of food patches occurred in a step-by-step manner, one patch attracting animals when a nearby patch had been depleted, and not following a model of ideal free distribution.Although our experimental food patches were exploited in relation to their distance from shelter, we were able to demonstrate that distance did not influence the dynamics of exploitation of a food item. The mean number of cockroaches on a food patch, whatever its spatial position, increased regularly, reached a maximum at t=–10 min, and then decreased rapidly after all the food had been completely consumed, at t=0 min. The mean number of animals in the 20 cm diameter circle round a food source peaked at t=0 min, then decreased rapidly. This area appeared to be a transit area. The mean number of animals in a 60 cm diameter circle round the food source peaked later, and then decreased slowly. Animals remained in this area longer than in the area closer to the food dish, but their presence there was concomitant with the depletion of the food box.     

Feeding rate as valuable information in primate feeding ecology

In this review I outline studies on wild non-human primates using information on feeding rate, which is defined as the food intake per minute on a dry-weight basis; further, I summarize the significance of feeding rate in primate feeding ecology. The optimal foraging theory has addressed three aspects of animal feeding: (1) optimal food patch choice, (2) optimal time allocation to different patches, and (3) optimal food choice. In order to gain a better understanding of these three aspects, the feeding rate itself or its relevance indices (e.g., rates of calorie and protein intake) could be appropriate measures to assess the quality of food and food patches. Moreover, the feeding rate plays an essential role in estimation of total food intake, because it varies greatly for different food items and the feeding time is not a precise measure. The feeding rate could also vary across individuals who simultaneously feed on the same food items in the same food patch. Body size-dependent and rank-dependent differences in the feeding rate sometimes cause individuals to take strategic behavioral options. In the closing remarks, I discuss the usefulness of even limited data on feeding rate obtained under adverse observational conditions in understanding primate feeding ecology.     

Living in forest fragments reduces group cohesion in diademed sifakas (Propithecus diadema) in eastern Madagascar by reducing food patch size

Forest fragmentation is thought to threaten primate populations, yet the mechanisms by which this occurs remain largely unknown. However, fragmentation is known to cause dietary shifts in several primate species, and links between food resource distribution and within-group spatial dynamics are well documented. Thus, fragmentation has the potential to indirectly affect spatial dynamics, and these changes may present additional stresses to fragmented populations. I present the results from a 12-month study of Propithecus diadema at Tsinjoarivo, eastern Madagascar, including two groups in fragments and two in continuous forest. Instantaneous data on activity and spatial position were collected during all-day focal animal follows. Fragment groups had much lower cohesion, being more likely to have no neighbor within 5 and 10 m. For continuous forest groups, cohesion was highest in the rainy season (when food patches are large) and lowest in winter (when the animals rely on small-crowned mistletoes), and the chance of having no neighbor within 5 m was positively correlated with mistletoe consumption. Thus their decreased cohesion in fragment groups is inferred to result from their increased reliance on mistletoes and other small resources, which causes them to spread out among multiple patches. This scenario is consistent with the reduced body mass of subordinate individuals (males and immatures) in fragments, and suggests the occurrence of steeper within-group fitness gradients. Further research is necessary to determine whether these patterns apply to other primates; however, since fragmentation tends to cause the loss of the largest trees, many primates in fragments may lose their largest food resources and undergo similar behavioral shifts.     

Within-group female-female agonistic interactions in Taiwanese macaques (Macaca cyclopis)

Feeding-related agonism among wild female Taiwanese macaques was investigated in two study groups at ecologically diverse sites (Fu-shan and Ken-ting) to determine whether contest-feeding competition was present in these groups. Females that contest for food within a primate group are hypothesized to form dominance hierarchies and tend to be philopatric. In this study we tested 1) whether Taiwanese macaque females show higher agonism in a feeding context, 2) whether they exhibit stronger agonism over higher-quality foods, and 3) whether higher agonism rates occur in smaller food patches. Female Taiwanese macaques at both study sites showed similar agonism rates in a feeding context (0.30 events/hr). They exhibited higher agonism rates inside food patches than outside food patches in the spring. Higher agonism rates occurred during seasons of higher fruit availability, and a lower agonism rate occurred in winter when the macaques switched to feeding on fallback foods. Females in the Fu-shan group exhibited higher proportions of aggressive interactions over higher-quality foods, such as animal matter and the reproductive parts of plants. In the Ken-ting group, 95.8% of feeding-related agonistic interactions among females occurred over fruits. Agonistic interactions that occurred in small food patches tended to result in the agonism recipient leaving the food patch. We conclude that female Taiwanese macaques show contest feeding competition in certain contexts. The patterns we observed have also been documented in other primate species in which females are philopatric and form linear dominance hierarchies.     

The effect of depletion and predictability of distinct food patches on the timing of aggression in red deer stags

For group-foraging ungulates, forage is generally widely and relatively evenly dispersed. However, for free-ranging red deer Cervus elaphus supplementary winter feeding provides distinct patches of predictable food. These patches differ in size, but also in temporal distribution and depletion rate. Interference competition is known to increase with increasing spatial clumping (decreasing patch size), but the influence of temporal clumping and the predictability of food occurrence has received much less attention. Therefore in this study we investigated the effects of different degrees of spatial and temporal clumping of food on interference competition during feeding. Patch size was the main parameter influencing participation in feeding as well as interference competition during feeding on the respective patch. Temporal dispersion and the predictability of food occurrence were however, important parameters for the timing of aggressive interactions. Generally, aggression occurs during feeding and increases with decreasing patch size. But when depletion rate was high food availability was predictably short and the patch occurred predictably (such as hay), middle ranking stags increased aggression already prior to feeding at the respective patch. We suggest that in this way they confirmed hierarchy outside feeding on the quickly depleted patch and as a result gained actual feeding time when feeding on the respective patch. With the patch occurring predictably but varying in size the number of participating subordinates varied concomitantly with variation in patch size. Subordinates assessed patch profitability and left without having fed when patch size was too small for an efficient participation. When patch size was predictably small enough to be defended exclusively (feed blocks), subordinate stags did not assess profitability each time but did not participate at all in feeding at the respective patch. The relative importance of these various food-related parameters (patch size depletion rate, predictability) influencing feeding competition and the timing of aggression will vary with group size, rank, alternative food sources, physical characteristics of the food as well as different hierarchy systems of the feeding animals.     

Anticipation of conflict by chimpanzees

Captive chimpanzees appear to anticipate the occurrence of conflict during feeding by grooming and being in proximity at increased rates during the hour prior to feeding. The effect is more marked when food is clumped than when it is dispersed, suggesting that the proximate cause is the anticipation of increased levels of competition. Chimpanzees did not choose high ranking individuals more often as prefeed grooming partners; rather, they preferred to associate with their normal grooming partners (as reflected in post-feed grooming preferences) and close kin. A strong correlation between prefeed association patterns and spatial proximity during clumped feeding sessions suggests that their main concern is to be allowed to feed near individuals who are able to monopolize food sources.     

Influences of sex,group size,and spatial position on vigilance behavior of Przewalski’s gazelles

Group-living animals may need to spend less time being vigilant, consequently, having more time for other important activities such as foraging (i.e., group size effect). Przewalski’s gazelle (Procapra przewalskii) is a group-living social animal, and a study was conducted in Qinghai Province of China during June–August 2006 by using a continuous focal sampling method to investigate the influences of group size, sex, within-group spatial position, and nearest-neighbor distance on individual vigilance level (defined as scanning frequency per minute). Male gazelles were more vigilant than females. The gazelle’s vigilance level decreased with group size (group size effect), but only for females. The individuals at the central positions within a group were less vigilant than those at the peripheral positions, but the nearest-neighbor distance did not have any significant influence on the individual vigilance level. Our results support the hypotheses of group size effect and edge effects, but the sexual difference in vigilance level and in the response to group size effect on vigilance suggests that there may be sexual difference in the function and targets of vigilance behavior of Przewalski’s gazelles, which warrants more investigation, with incorporation of within-group spatial position, to better understand the mechanism underlying the group size effect and edge effect.     

Tolerance of understory plants subject to herbivory by roe deer

Classical foraging theory states that animals feeding in a patchy environment can maximise their long term prey capture rates by quitting food patches when they have depleted prey to a certain threshold level. Theory suggests that social foragers may be better able to do this if all individuals in a group have access to the prey capture information of all other group members. This will allow all foragers to make a more accurate estimation of the patch quality over time and hence enable them to quit patches closer to the optimal prey threshold level. We develop a model to examine the foraging efficiency of three strategies that could be used by a cohesive foraging group to initiate quitting a patch, where foragers do not use such information, and compare these with a fourth strategy in which foragers use public information of all prey capture events made by the group. We carried out simulations in six different prey environments, in which we varied the mean number of prey per patch and the variance of prey number between patches. Groups sharing public information were able to consistently quit patches close to the optimal prey threshold level, and obtained constant prey capture rates, in groups of all sizes. In contrast all groups not sharing public information quit patches progressively earlier than the optimal prey threshold value, and experienced decreasing prey capture rates, as group size increased. This is more apparent as the variance in prey number between patches increases. Thus in a patchy environment, where uncertainty is high, although public information use does not increase the foraging efficiency of groups over that of a lone forager, it certainly offers benefits over groups which do not, and particularly where group size is large.     

Food Supply and Chimpanzee (Pan troglodytes schweinfurthii) Party Size in the Budongo Forest Reserve,Uganda

A central issue in socioecology is the nature of the relationship between an organism's environment and its social structure. In chimpanzees, the fission-fusion social system is thought to minimize feeding competition for primary dietary components: ephemeral, dispersed patches of ripe fruit. Intragroup feeding competition is thought to force individuals into small parties. Informal observations in the Sonso region of the Budongo forest had suggested that in this habitat, food supply was such that feeding competition was less important in determining grouping patterns than elsewhere. We used data collected on food supply and party sizes over a 4-year period to investigate this suggestion. In accord with theoretical expectation, sizes of foraging parties fluctuated with the size of food patches. However, party sizes showed either negative or no relationship with habitat-wide measures of food abundance. Likewise party sizes showed little relationship to overall measures of food dispersion. For important dietary items, both fruit and leaves had patchy distributions, though the degree of clumping was not strong, and fruit was not more clumped than leaves. Generally, abundant food appeared to be less patchy, and chimpanzees appeared to use more patches as food became more abundant rather than forming larger parties. We suggest that both dispersal and abundance need to be considered when investigating the impact of food supply on grouping patterns, and that the importance of food as a factor in determining chimpanzee grouping patterns declines with increasing levels of abundance.     

营养状态和代谢范围对锦鲫幼鱼群体行为的影响

为考察营养状态和代谢范围对鱼类群体行为的影响,研究以锦鲫(Carassius auratus)幼鱼为实验对象,在(25.4±0.2)℃条件下先测定其摄食代谢和能量代谢(标准代谢率, SMR;最大代谢率, MMR)计算代谢范围(AS=MMR–SMR),再测定5个“营养-AS”处理组的锦鲫鱼群体中的个体空间位置、摄食量及个体特征(如个体游泳速度和加速度)和群体特征(如个体游泳速度同步性、个体间距离、最近邻距离和群体极性)。研究发现:营养状态、饥饿、代谢范围、摄食和消化对鱼群中的个体空间位置均无影响。饥饿和消化对锦鲫群体的凝聚力并无影响,但饥饿降低该种鱼群体协调性的现象仅在消化期间存在,即群体中个体食物获取能力导致消化策略并非相同,由此引发个体游泳运动同步性更加紊乱,最终导致群体协调性下降。在正常营养状态的锦鲫群体中,群体前部的空间可赋予个体获得更多食物资源的生态收益,但饥饿消除该群体中个体空间分布生态收益的异质性。对照组摄食量与摄食水平与预测剩余的AS呈负相关,饥饿组摄食量与摄食水平与预测剩余的AS不相关。研究表明:在正常营养状态的锦鲫群体中,群体前部的空间可赋予个体获得更多食物资源的生态...     

Quantifying Primate Food Distribution and Abundance for Socioecological Studies: An Objective Consumer-centered Method

Food abundance and distribution have played a central role in the conceptual theory of primate socioecology. 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, nonagonistic (scramble) competition is expected. Primatologists usually measure resource density and patchiness from a botanical perspective, without an explicit link to the biology of the animal being studied. Such an approach may be irrelevant to how the animals view the dispersion of resources. For studies related to feeding competition, we suggest the use of a method that provides a consumer-based index of food distribution. We then describe such an approach and apply it to understand agonistic behavior in white-faced capuchins (Cebus capucinus), at Lomas Barbudal. Instead of choosing sample plots at random, we use each actual feeding tree of a group as the center of a sample plot and we use the monkey species’ average group spread as the sampling area. This focal tree method allows us to evaluate the resource availability both within and outside of the feeding tree during a particular feeding bout. To summarize the spatial distribution of food at the level of a foraging group, we define and use an extension of Lloyd’s Dispersion Index, Lloyd’s Extended Index (LEI), designed to allow the inclusion of resources of diverse sizes and species in a single measure. We evaluate if LEI can be used to predict the frequency of aggression, if changing the area of the plot alters these results, and if calculating LEI based on fruit abundance or fruit biomass better predicts the frequency of aggression in this population of capuchins. In support of socioecological predictions, our results show that the frequency of agonism in a focal tree declines as LEI increases. This relationship is significant when LEI is calculated using a 20-m plot size and weighting tree size by fruit counts, but not when using larger plot sizes, unweighted tree counts, or weighting by fruit biomass. Our approach demonstrates the importance of carefully considering plot size and different measures of food availability when testing socioecological models relating resource distribution and quality to aggression in nonhuman primates.     

Group foraging in wild brown hares: effects of resource distribution and social status

Wild brown hares (Lepus europaeus), though normally comparatively solitary, have the capacity to adjust their behaviour such that they can benefit when foraging in groups. They are able to allocate more time to feeding and have an increased corporate vigilance as group size increases. However, these benefits are conditional upon the food distribution. When food is spaced, all individuals benefit. When it is clumped into a small defendable patch, dominant hares attempt to monopolize the resource. They can successfully exclude subordinates when group size is small but, as group size increases, they must devote more time to defending the patch. Therefore, dominants spend less time feeding with increasing group size, while subordinates spend more, since they have more opportunity to feed while the dominant hare is off chasing other individuals. As a consequence, when more than two hares are present, all individuals do less well when food is clumped than when it is spaced.     

Optimal foraging theory predicts effects of environmental enrichment in a group of adult golden lion tamarins

The success of environmental enrichment programs in effecting specific changes in the behavior of captive animals has not always been uniform. Separate studies demonstrated both an increase in food competition and a decrease in food competition among captive group-living primates upon introduction of foraging devices. The objectives of this study were to measure the effects of variation in resource distribution and availability on food competition in a group of captive adult golden lion tamarins (Leontopithecus rosalia). The resource variables chosen were suggested from optimal foraging theory. The energy invested to obtain an item influenced food transfer and aggressive behaviors while food abundance did not. All individuals obtained an equivalent number of items over the course of the experiment from the foraging device, even though some tamarins obtained most of their food rewards directly from the device while others received their food rewards primarily through food transfer from other group members. Because the monkeys appeared highly motivated to obtain food from the test apparatus and did not habituate to it, the foraging device used in this experiment could be used as regular environmental enrichment for golden lion tamarins. One way to circumvent potentially unacceptable rates of aggression, with this or any feeding protocol that increases foraging task complexity and search time, may be to provide more than one foraging device per group. Zoo Biol 17:231–244, 1998. © 1998 Wiley-Liss, Inc.     

Dominance competition through affiliation and support in Japanese macaques: An experimental study

It has been proposed that monkeys direct grooming to high-ranking individuals in an attempt to obtain agonistic support in return. But whether these two categories of interactions are causally related has proven difficult to establish. Part of the problem stems from the fact that in stable groups social relationships reflect an equilibrium state and that behaviors need only be performed at low rates and long intervals to maintain the current social structure. In theory, however, if affiliative and supportive interactions are indeed causally related, it should be possible to accentuate their temporal relation, hence their causal dynamics. For example, destabilizing dominance relations can be expected to induce competition for status and force individuals to deploy behavioral tactics for settling new rank relations. We experimentally induced rank reversals in a captive group of Japanese macaques (Macaca fuscata) composed of three matrilines (A-B-C rank order). A reversed C-A-B order composed of three individuals per matriline was maintained for 2 weeks. The results show the close temporal relation among (i) asserting one’s rank, (ii) competing for access to dominants through affiliation and interferences in affiliation, (iii) receiving support from dominants against lower-ranking individuals, and (iv) supporting dominants against subordinates. These findings are compatible with one version of the affiliation-for-support hypothesis, namely that monkeys affiliate with dominants as a way to assert their position in the hierarchy. In a functional perspective, mutual selfishness provides a better explanation than reciprocal altruism because the possibility that both groomers and supporters derive immediate net benefits cannot be excluded.     

Food patch structure and plant resource partitioning in interspecific associations of amazonian tamarins

ávila-Pires’ saddle-back tamarins (Saguinus fuscicollis avilapiresi) and red-cap moustached tamarins (S. mystax pileatus), coexisting in highly stable mixed-species groups, overlapped considerably in their use of plant food resources at an Amazonian terra firme forest site. Overlap between food types consumed by the two species was particularly high during periods of lowest fruit availability, when they resorted to a common food supply, primarily the pod exudates of two emergent species of legume trees (Parkia nitida andParkia pendula) and nectar ofSymphonia globulifera. Within-group interspecific competition did not covary with independent measures of resource availability, contrary to predictions based on resource partitioning models. A greater number of both saddle-back and moustached tamarins were able to feed for longer patch residence periods within larger and more productive food patches, whereas small and clumped patches could be monopolized by the socially and numerically dominant moustached tamarins to the physical exclusion of the smaller-bodied saddle-back tamarins. Overall rates of interspecific aggression were extremely low, however, partly because patches that could be monopolized contributed with a minor proportion of either species’ diet. Saddle-backs foraged at lower levels in the understory and encountered smaller food patches more often, whereas moustached tamarins foraged higher and encountered more larger patches in the middle canopy. Although the two species led one another to differently-sized patches, moustached tamarins initiated most feeding bouts and encountered significantly larger and more productive patches that tended to accommodate the entire mixed-species group. Disadvantages of exploitative and interference feeding competition over plant resources, and advantages of shared knowledge of food patches, are but one component of the overall cost-benefit relationship of interspecific associations in tamarins.