Predation hazard and seed removal by small mammals: microhabitat versus patch scale effects

Predator avoidance may involve response strategies of prey species that are time and space specific. Many studies have shown that foraging individuals avoid predators by altering microhabitat usage; alternatively, sites may be selected according to larger-scale features of the habitat mosaic. We measured seed removal by two small mammal species (Peromyscus leucopus, and Microtus pennsylvanicus) at 474 stations over an experimentally created landscape of 12 patches, and under conditions of relatively high (full moon) and low (new moon) predatory hazard. Our objective was to determine whether predator avoidance involved the selection of small-, medium-, or large-scale features of the landscape (i.e., at the scale of microhabitats, habitats, or habitat patches). We found rates of seed removal to vary more with features of whole patches than according to variation in structural microhabitats within patches. Specific responses included: under-utilization of patch edge habitats during full moon periods, and microhabitat effects that were only significant when considered in conjunction with larger-scale features of the landscape. Individuals residing on large patches altered use of microhabitats/habitats to a greater extent than those on smaller patches. Studies just focusing on patterns of microhabitat use will miss responses at the larger scales, and may underestimate the importance of predation to animal foraging behavior.     

Microhabitat use by foraging white-clawed crayfish (<Emphasis Type="Italic">Austropotamobius pallipes</Emphasis>) in stream pools in the NE Iberian Peninsula

The white-clawed crayfish (Austropotamobius pallipes) is an endangered species across most of its distribution range, and information on its ecological requirements is needed to implement effective conservation measures. Its habitat use has been studied in different areas and at various spatial scales. However, being a nocturnal species, there is scarce information on its habitat selection during foraging periods. In this work we analyse nocturnal habitat use of white-clawed crayfish in pools of a small stream in the northeastern Iberian Peninsula at two different scales: (1) microhabitat selection and (2) pool characteristics. Large crayfish showed a clear positive selection for deeper microhabitats, a selection pattern that was weaker for medium-sized crayfish and absent for small ones. On the other hand, crayfish of all sizes avoided cobble and boulder microhabitats and positively selected fine substrate and more exposed microhabitats. Crayfish abundance in pools was positively influenced by pool area, pool depth and the availability of fine substrates, especially silt. While studies on white-clawed crayfish habitat use have often stressed the importance of rough substrates as crayfish refuge, our results show that fine substrates are positively selected by foraging crayfish of all size classes and promoted active crayfish abundance in pools. These apparently contradictory results may be due to the differences in microhabitat preferences exhibited by active and inactive crayfish. Thus, our results help to better complete the picture of white-clawed crayfish habitat requirements.     

Feeding guild structure of beetles on Australian tropical rainforest trees reflects microhabitat resource availability

1.?We tested the hypotheses that feeding guild structure of beetle assemblages changed with different arboreal microhabitats and that these differences were consistent across rainforest tree species. 2.?Hand collection and beating techniques were used from the gondola of the Australian Canopy Crane to collect beetles from five microhabitats (mature leaves, flush leaves, flowers, fruit and suspended dead wood) within the rainforest canopy. A simple randomization procedure was implemented to test whether the abundances of each feeding guild on each microhabitat were different from that expected based on a null hypothesis of random distribution of individuals across microhabitats. 3.?Beetles from different feeding guilds were not randomly distributed, but congregated on those microhabitats that are likely to provide the highest concentrations of their preferred food sources. Herbivorous beetles, in particular, were over-represented on flowers and flush foliage and under-represented on mature leaves and dead wood. Proportional numbers of species within each feeding guild were remarkably uniform across tree species for each microhabitat, but proportional abundances of feeding guilds were all significantly non-uniformly distributed between host tree species, regardless of microhabitat, confirming patterns previously found for arthropods in trees in temperate and tropical forests. 4.?These results show that the canopy beetle community is partitioned into discrete assemblages between microhabitats and that this partitioning arises because of differences in feeding guild structure as a function of the diversity and the temporal and spatial availability of resources found on each microhabitat.     

Exploring the nature of ecological specialization in a coral reef fish community: morphology,diet and foraging microhabitat use

Patterns of ecological specialization offer invaluable information about ecosystems. Yet, specialization is rarely quantified across several ecological niche axes and variables beyond the link between morphological and dietary specialization have received little attention. Here, we provide a quantitative evaluation of ecological specialization in a coral reef fish assemblage (f. Acanthuridae) along one fundamental and two realized niche axes. Specifically, we examined ecological specialization in 10 surgeonfish species with regards to morphology and two realized niche axes associated with diet and foraging microhabitat utilization using a recently developed multidimensional framework. We then investigated the potential relationships between morphological and behavioural specialization. These relationships differed markedly from the traditional ecomorphological paradigm. While morphological specialization showed no relationship with dietary specialization, it exhibited a strong relationship with foraging microhabitat specialization. However, this relationship was inverted: species with specialized morphologies were microhabitat generalists, whereas generalized morphotypes were microhabitat specialists. Interestingly, this mirrors relationships found in plant–pollinator communities and may also be applicable to other ecosystems, highlighting the potential importance of including niche axes beyond dietary specialization into ecomorphological frameworks. On coral reefs, it appears that morphotypes commonly perceived as most generalized may, in fact, be specialized in exploiting flat and easily accessible microhabitats.     

Effects of body size,seed density,and soil characteristics on rates of seed harvest by heteromyid rodents

Summary Although it is well established that coexisting heteromyid rodent species forage in different microhabitats, we do not yet know the basis for divergent microhabitat choice. One possibility is that seed harvest rates differ among microhabitats, and each species forages where it can extract seeds most efficiently. Microhabitats vary in several factors that could affect heteromyid foraging efficiency, including seed density, soil organic content and particle size distribution. We have explored the effect of each of these variables on harvest rates of several species feeding from petri dishes containing known densities of millet seeds embedded in soil of known particle size and density. Results indicate that the number of seeds harvested per second increases uniformly with seed density and soil density and decreases with soil particle size. Body size affects these relationships: larger animals have higher harvest rates for a given set of conditions and experience a greater relative change in harvest rate for a give change in conditions. This implies that heteromyids can be expected to exhibit species-specific microhabitat preferences while foraging in nature.     

Foraging strategies of an insectivorous marsupial,Sminthopsis youngsoni (Marsupialia: Dasyuridae), in Australian sandridge desert

This study investigated the effects of predation risk, dune position and microhabitat on foraging of the Lesser Hairy‐footed Dunnart Sminthopsis youngsoni, a small insectivorous marsupial, in the Simpson Desert of western Queensland. The intensity of foraging was assessed by establishing feeding stations (dishes containing mealworms) in open and bush microhabitats at three levels on sand dunes, and recording the numbers of prey taken by dunnarts from the stations after nightly bouts of foraging. Risk of predation was manipulated by provision of artificial illumination at alternate feeding stations on each of five occasions when trials were run. The numbers of mealworms left after feeding bouts varied inconsistently across trials, providing little evidence that dunnarts respond to habitat or predation risk while foraging. These results contrast sharply with studies of arid zone rodents, where foraging is usually sensitive to both predation risk and resource distribution. We suggest that S. youngsoni forages equally in all habitats of its sandridge environment, and experiences relatively low risk of predation whilst doing so.     

Exploration and exploitation of foraging patches by desert sparrows: environmental indicators and local evaluation of spatially correlated costs and benefits

Conventional evolutionary and behavioral reasoning expects foragers to show strong spatial preferences in environments with heterogeneous resource distribution. Moreover, consumers should benefit from exploiting the information embedded in environmental features that indicate resource abundance. In desert soils seed abundance associates strong and reliably with vegetation and litter cover at small spatial scales. However, other spatially correlated factors (substrate complexity, temperature, predation risk) may affect foraging costs, benefits and decisions by ground‐feeding granivores. We used a sequence of three semi‐controlled field experiments of binary spatial choice within a portable aviary to identify the main cause of foraging microhabitat selection by the most abundant postdispersal granivorous bird in the central Monte desert (Argentina). In the first experiment we placed the aviary at field to offer pairs of adjacent microhabitats of unmodified, naturally‐contrasting substrates and environmental conditions to single, untrained rufous‐collared sparrows Zonotrichia capensis. Birds selected covered microhabitats in winter and summer, ruling out substrate complexity or thermoregulation as main single causes of patch selection. The other two experiments dissociated seed abundance, tree cover and litter to reveal their effects on patch selection. The results indicate that 1) sparrows do not restrict microhabitat exploration relying on environmental indicators, 2) distance to tree cover influences the order of patch exploration, probably in association with apprehension or risk‐assessment behavior, and 3) patch exploitation is determined by short‐term local estimation of seed abundance. The integration of these with previous results obtained under variable degrees of realism and experimental control allows for a better explanation of the spatial component of postdispersal granivory and its consequences on plants. The unconstrained selective foraging strategy of these sparrows would allow them to detect sporadic or ephemeral rich patches with structural characteristics indicating ‘low‐quality’, should promote the spatial homogenization of the palatable seed bank, and would favor indirect interactions between plants.     

Multi-Scale Foraging Habitat Use and Interactions by Sympatric Cervids in Northeastern China

Abstract: Moose (Alces alces) and roe deer (Capreolus pygargus) are sympatric in the forest region of northeastern China. Using univariate analyses of feeding sign data, we found the 2 species were positively associated, but there were distinctions between their use of forage resources across landscape, patch, and microhabitat scales. We used resource selection function models to predict the influence of environmental covariates on moose and roe deer foraging; we detected covariate effects at the landscape and microhabitat scales but not at the patch scale. Forage resources used by the 2 species were similar, but moose used wetter areas and more low-visibility habitats than did roe deer, which strongly avoided areas with sparse vegetation. Both species were influenced by forage abundance and distribution at the microhabitat scale but exhibited differences in intensity of use of plant species and microhabitats. Moose used areas with deeper snow and avoided hiding cover; roe deer avoided areas with higher total basal areas of tree stems and preferred areas with high plant species richness. For moose, there was a trade-off in the use of concealment cover between the landscape and microhabitat scales. We detected avoidance by moose of roads where roe deer occurred. Roe deer exhibited more capacity for coping with human disturbance and interspecific interaction. In areas similar to our study area, road closures and suppression of roe deer near roads within 3–5 years postlogging may benefit moose. Furthermore, a mosaic of areas with different logging intervals may contribute to spatial separation of moose and roe deer and promote their coexistence.     

How does the presence of a conspecific individual change the behavioral game that a predator plays with its prey?

Behavioral games predators play among themselves may have profound effects on behavioral games predators play with their prey. We studied the behavioral game between predators and prey within the framework of social foraging among predators. We tested how conspecific interactions among predators (little egret) change the predator–prey behavioral game and foraging success. To do so, we examined foraging behavior of egrets alone and in pairs (male and female) in a specially designed aviary consisting of three equally spaced pools with identical initial prey (comet goldfish) densities. Each pool was comprised of a risky microhabitat, rich with food, and a safe microhabitat with no food, forcing the fish to trade off food and safety. When faced with two versus one egret, we found that fish significantly reduced activity in the risky habitat. Egrets in pairs suffered reduced foraging success (negative intraspecific density dependence) and responded to fish behavior and to their conspecific by changing their visiting regime at the different pools—having shorter, more frequent visits. The time egret spent on each visit allowed them to match their long-term capture success rate across the environment to their capture success rate in the pool, which satisfies one aspect of optimality. Overall, egrets in pairs allocated more time for foraging and changed their foraging tactics to focus more on fish under cover and fish ‘peeping’ out from their shelter. These results suggest that both prey and predator show behavioral flexibility and can adjust to changing conditions as needed in this foraging game.     

Predator avoidance,microhabitat shift,and risk-sensitive foraging in larval dragonflies

Summary Dragonfly larvae (Odonata: Anisoptera) are often abundant in shallow freshwater habitats and frequently co-occur with predatory fish, but there is evidence that they are underutilized as prey. This suggests that species which successfully coexist with fish may exhibit behaviors that minimize their risk of predation. I conducted field and laboratory experiments to determine whether: 1) dragonfly larvae actively avoid fish, 2) microhabitat use and foraging success of larvae are sensitive to predation risk, and 3) vulnerability of larvae is correlated with microhabitat use. I experimentally manipulated the presence of adult bluegills (Lepomis macrochirus) in defaunated patches of littoral substrate in a small pond to test whether colonizing dragonfly larvae would avoid patches containing fish. The two dominant anisopteran species, Tetragoneuria cynosura and Ladona deplanata (Odonata: Libellulidae), both strongly avoided colonizing patches where adult bluegills were present. Laboratory experiments examined the effects of diel period and bluegills on microhabitat use and foraging success, using Tetragoneuria, Ladona and confamilial Sympetrum semicictum, found in a nearby fishless pond. Tetragoneuria and Ladona generally occupied microhabitats offering cover, whereas Sympetrum usually occupied exposed locations. Bluegills induced increased use of cover in all three species, and use of cover also tended to be higher during the day than at night. Bluegills depressed foraging in Tetragoneuria and to a lesser extent in Ladona, but foraging in Sympetrum appeared unaffected. Other laboratory experiments indicated that Sympetrum were generally more vulnerable than Tetragoneuria or Ladona to bluegill predation, and that vulnerability was positively correlated with use of exposed microhabitats. Both fixed (generally low use of exposed microhabitats, diel microhabitat shifts) and reactive (predator avoidance, predator-sensitive microhabitat shifts) behavioral responses appear to reduce risk of predation in dragonfly larvae. Evidence indicates that vulnerability probably varies widely among species and even among instars within species, and suggests that spatial distributions of relatively vulnerable species may be limited by their inability to avoid predation.     

Growth-mortality tradeoffs and 'personality traits' in animals

Consistent individual differences in boldness, reactivity, aggressiveness, and other 'personality traits' in animals are stable within individuals but vary across individuals, for reasons which are currently obscure. Here, I suggest that consistent individual differences in growth rates encourage consistent individual differences in behavior patterns that contribute to growth-mortality tradeoffs. This hypothesis predicts that behavior patterns that increase both growth and mortality rates (e.g. foraging under predation risk, aggressive defense of feeding territories) will be positively correlated with one another across individuals, that selection for high growth rates will increase mean levels of potentially risky behavior across populations, and that within populations, faster-growing individuals will take more risks in foraging contexts than slower-growing individuals. Tentative empirical support for these predictions suggests that a growth-mortality perspective may help explain some of the consistent individual differences in behavioral traits that have been reported in fish, amphibians, reptiles, and other animals with indeterminate growth.     

Microhabitat partitioning in seagrass mesograzers is driven by consistent species choices across multiple predator and competitor contexts

Explanations for the coexistence of multiple species from the same functional group or taxonomic clade frequently include fine‐scale resource partitioning. However, despite the hypothesized importance of niche partitioning, we know relatively little about the underlying mechanisms. For example, differences in resource use may be fixed consequences of organism traits, or they may be achieved via context‐dependent behaviors. In this study we investigated mechanisms of microhabitat partitioning using eight species of marine mesograzers inhabiting seagrass and algae habitats, using laboratory trials to measure microhabitat use in the presence and absence of both predators and competitors. We found clear evidence for microhabitat partitioning between the species, which account for over 60% of the mesograzers commonly found in this system and vary in both body size and the ability to build tubes on habitat substrates. Species‐specific microhabitat use was poorly predicted by these two traits, but remained remarkably consistent across contexts. Habitat use was not affected by the presence of fish predators common in this system, even though predation pressure is thought to place strong constraints on microhabitat in communities of plant‐associated arthropods. The presence of competing species also did not affect the relative separation of microhabitat use. Behavioral responses to potential competitors did cause significant changes in microhabitat use in all of the smallest species, but these changes did not depend on competitor identity and were relatively small compared to among‐species patterns of microhabitat partitioning. The consistency of species‐specific microhabitat use, regardless of the presence of predators or competitors, should make coexistence most likely among species that differ in these choices. For these species, it appears that the benefits accrued from their selected microhabitats are not affected by species interactions, or that any benefits of alternative microhabitat use are outweighed by risks associated with movement.     

Food resource matching by foraging tits Parus spp. during spring-summer in a Mediterranean mixed forest; evidence for an ideal free distribution

We analysed whether patterns of microhabitat use by Blue Tits Parus caeruleus , Great Tits Parus major and Crested Tits Parus cristatus inhabiting a mixed forest consistently matched the patterns of food availability experienced by foraging birds during spring-summer. The use of five microhabitats by each bird species (the foliage of three tree species, shrubs and ground) and the availability of food in trees during the prebreeding, breeding and post-breeding periods of the birds' annual cycle were measured. All three tit species foraged mainly in the outer part of tree canopies (small branches and leaves or needles). Tit distributions between tree species matched food resource distributions irrespective of overall food resource levels, which varied four-fold between the study periods, and tit species. Tits also exploited secondary microhabitats (shrubs and ground) in periods of low food availability; Blue Tits tended to use shrubs, whereas Great and Crested Tits foraged on the ground. Between-trees distributions fitted that expected from an ideal free distribution, suggesting that food availability and intraspecific exploitative competition were the main factors governing tree use by tits. In contrast, patterns of use of secondary microhabitats (shrubs and ground) seemed to indicate a role for the species-specific morphological configurations of each tit species since Blue Tits are better adapted to hang and tended to forage in shubs, whereas Great and Crested Tits are better adapted to feed on horizontal surfaces and tended to forage on the ground. No evidence of interspecific interactions was observed. Overall, the results pointed to an independent exploitation of Mediterranean mixed forest by each bird species, food availability and food accessibility being the main factors affecting microhabitat use by foraging tits.     

Non‐native trout limit native brook trout access to space and thermal refugia in a restored large‐river system

We used direct observation via snorkeling surveys to quantify microhabitat use by native brook (Salvelinus fontinalis) and non‐native brown (Salmo trutta) and rainbow (Onchorynchus mykiss) trout occupying natural and restored pool habitats within a large, high‐elevation Appalachian river, United States. Permutational multivariate analysis of variance (PERMANOVA) and subsequent two‐way analysis of variance (ANOVA) indicated a significant difference in microhabitat use by brook and non‐native trout within restored pools. We also detected a significant difference in microhabitat use by brook trout occupying pools in allopatry versus those occupying pools in sympatry with non‐native trout—a pattern that appears to be modulated by size. Smaller brook trout often occupied pools in the absence of non‐native species, where they used shallower and faster focal habitats. Larger brook trout occupied pools with, and utilized similar focal habitats (i.e. deeper, slower velocity) as, non‐native trout. Non‐native trout consistently occupied more thermally suitable microhabitats closer to cover as compared to brook trout, including the use of thermal refugia (i.e. ambient–focal temperature >2°C). These results suggest that non‐native trout influence brook trout use of restored habitats by: (1) displacing smaller brook trout from restored pools, and (2) displacing small and large brook trout from optimal microhabitats (cooler, deeper, and lower velocity). Consequently, benefits of habitat restoration in large rivers may only be fully realized by brook trout in the absence of non‐native species. Future research within this and other large river systems should characterize brook trout response to stream restoration following removal of non‐native species.     

Seasonal variation of tadpole spatial niches in permanent streams: The roles of predation risk and microhabitat availability

Phenotypic plasticity can improve fitness in unstable environments and can be expressed in many traits, such as life history attributes, growth and behavioural features. Microhabitat choice can have important consequences for development and survival of aquatic organisms and is expected to vary in response to stimuli, such as predation risk, food availability and temperature. At seasonal sites, microhabitat availability and associated benefits may change from season to season, which might lead to altered patterns of microhabitat use by tadpoles. We investigated this hypothesis in 17 streams from two localities in south‐eastern Brazil. We tested whether water level drops significantly during the dry season, whether lower water level results in altered microhabitat availability and whether predation risk changes between seasons, based on predator density. We then tested whether tadpoles change their pattern of microhabitat use, their spatial niche breadth (given by diversity of used microhabitats) and spatial niche overlap (in the case of co‐occurring species). We were able to include in our analyses tadpoles of four species of Hylidae, that occurred throughout both seasons. Stream depth decreased in the dry season, but microhabitat availability remained relatively stable in many streams, and predator density did not change significantly. Tadpoles of three out of the four species studied were more abundant during the dry season, which may be an adaptation to adjust time of metamorphosis to the rainy season. Tadpoles changed their patterns of microhabitat use between seasons, although the potential causing factors investigated did not seem to be responsible. Tadpole plasticity in microhabitat use may indicate the existence of selective pressures that vary through time and space and are still not well understood.     

Foraging behaviour of non-breeding Western Sandpipers Calidris mauri as a function of sex, habitat and flocking

Individuals within a population may vary considerably in the way they exploit available food resources. If the sexes differ in the size of their feeding apparatus, there can be differences in foraging behaviour and habitat use, hence one sex may be more susceptible to competition. We examined relationships between sexual dimorphism in bill size and foraging behaviour, and habitat and microhabitat use of non-breeding Western Sandpipers Calidris mauri at Bahía Santa María, northwestern Mexico. Western Sandpipers are sexually dimorphic, with females about 15% longer-billed than males. Males used a higher proportion of pecks, had a lower probing–pecking rate, walked at higher rates, foraged at sites with lower water content, and had greater variation in foraging technique than females. Moreover, males decreased their proportion of pecks and foraged at a higher rate than females when they changed from feeding alone to feeding in flocks, suggesting a greater safety advantage or susceptibility to conspecific interference when flock feeding. We compared behaviour and microhabitat usage in three habitats: brackish flats, mangroves, and cattail marshes. Sex-related differences in foraging behaviour and microhabitat use were consistent among habitats. Birds in brackish flats and mangroves used a higher proportion of pecks, foraged at lower rates and walked at higher rates, and foraged at deeper sites, with a lower proportion of water cover, than those in cattail marshes. Sex-related differences in foraging behaviour and microhabitat should reduce the level of competition between sexes, and may account for sex differences in Western Sandpiper distribution observed between habitats in Bahía Santa María.     

The foraging behavior of Japanese macaques Macaca fuscata in a forested enclosure:Effects of nutrient composition,energy and its seasonal variation on the consumption of natural plant foods

In the wild, primate foraging behaviors are related to the diversity and nutritional properties of food, which are affected by seasonal variation. The goal of environmental enrichment is to stimulate captive animals to exhibit similar foraging behavior of their wild counterparts, e.g. To extend foraging time. We conducted a 12-month study on the foraging behavior of Japanese macaques in a semi-naturally forested enclosure to understand how they use both provisioned foods and naturally available plant foods and what are the nutritional criteria of their consumption of natural plants. We recorded time spent feeding on provisioned and natural plant foods and collected the plant parts ingested of their major plant food species monthly, when available.We conducted nutritional analysis (crude protein, crude lipid, neutral detergent fiber-'NDF', ash) and calculated total non-slructural carbohydrate - 'TNC' and total energy of those food items. Monkeys spent 47% of their feeding time foraging on natural plant species. The consumption of plant parts varied significantly across seasons. We found that leaf items were consumed in months when crude protein, crude protein-to-NDF ratio, TNC and total energy were significantly higher and NDF was significantly lower, fruit/nut items in months when crude protein and TNC were significantly higher and crude lipid content was significantly lower, and bark items in months when TNC and total energy were higher and crude lipid content was lower. This preliminary investigation showed that the forested enclosure allowed troop members to more fully express their species typical flexible behavior by challenging them to adjust their foraging behavior to seasonal changes of plant item diversity and nutritional content, also providing the possibility for individuals to nutritionally enhance their diet.     

Peatland microhabitat heterogeneity drives phototrophic microbe distribution and photosynthetic activity

Phototrophic microbes are widespread in soils, but their contribution to soil carbon (C) uptake remains underexplored in most terrestrial systems, including C-accreting systems such as peatlands. Here, by means of metabarcoding and ecophysiological measurements, we examined how microbial photosynthesis and its biotic (e.g., phototrophic community structure, biomass) and abiotic drivers (e.g., Sphagnum moisture, light intensity) vary across peatland microhabitats. Using a natural gradient of microhabitat conditions from pool to forest, we show that the structure of phototrophic microbial communities shifted from a dominance of eukaryotes in pools to prokaryotes in forests. We identified five groups of co-occurring phototrophic operational taxonomic units with specific environmental preferences across the gradient. Along with such structural changes, we found that microbial C uptake was the highest in the driest and shadiest microhabitats. This study renews and improves current views on phototrophic microbes in peatlands, as the contribution of microbial photosynthesis to peatland C uptake has essentially been studied in wet microhabitats.     

Seasonal variation in the positional behavior of red howling monkeys (Alouatta seniculus)

Recent studies on the positional behavior of primates reveal that significant seasonal variation occurs in both locomotion and postures that is related to changes in diet and foraging techniques. Howling monkeys (genusAlouatta), which also have a seasonally varied diet, are predicted to have correspondingly varied positional behaviors. Two groups of red howling monkeys were studied in a primary rain forest in French Guinana during the dry and wet seasons. During the dry season, when howler diet is based mainly on leaves, howlers traveled more frequently by quadrupedal walking on large supports, a mode of progression that is probably inexpensive energetically and relatively stable. During feeding, quadrupedal and tripedal stand contributed considerably, a posture probably associated with the equal distribution of leaves within a tree crown. In contrast, during the wet season, when fruit was abundant, howlers fed very frequently by sitting on large supports, probably because fruit consumption required more time for special manipulation. However, most seasonal changes in feeding postures, and in travel and feeding locomotion, were difficult to associate directly with dietary shifts. These behavioral changes may be more highly correlated with slight modifications in microhabitat use (horizontal and vertical daily ranges, similar and alternative arboreal pathways) that are not considered in this paper.