Optimal foraging and community structure: implications for a guild of generalist grassland herbivores

Summary A particular linear programming model is constructed to predict the diets of each of 14 species of generalist herbivores at the National Bison Range, Montana. The herbivores have body masses ranging over seven orders of magnitude and belonging to two major taxa: insects and mammals. The linear programming model has three feeding constraints: digestive capacity, feeding time and energy requirements. A foraging strategy that maximizes daily energy intake agrees very well with the observed diets. Body size appears to be an underlying determinant of the foraging parameters leading to diet selection. Species that possess digestive capacity and feeding time constraints which approach each other in magnitude have the most generalized diets. The degree that the linear programming models change their diet predictions with a given percent change in parameter values (sensitivity) may reflect the observed ability of the species to vary their diets. In particular, the species which show the most diet variability are those whose diets tend to be balanced between monocots and dicots. The community-ecological parameters of herbivore body-size ranges and species number can possibly be related to foraging behavior.     

Body size,energetic and foraging mode of raptors in central Chile

Summary An inferential analysis of the foraging mode (opportunist or mediate by prey selection) of a taxonomic assemblage of raptors in central Chile was conducted. The analysis of energetic aspects such as daily requirements of predators and energy supplied by the preys, with estimations of prey relative abundances and their incidence in the diet of raptors, led us to conclude that contrarly to the opportunistic hunting mode suggested by others authors, these predators apparently present prey selection (on a biomass basis). This phenomenon is particularly evident in raptors of small body size.     

Bee foraging ranges and their relationship to body size

Bees are the most important pollinator taxon; therefore, understanding the scale at which they forage has important ecological implications and conservation applications. The foraging ranges for most bee species are unknown. Foraging distance information is critical for understanding the scale at which bee populations respond to the landscape, assessing the role of bee pollinators in affecting plant population structure, planning conservation strategies for plants, and designing bee habitat refugia that maintain pollination function for wild and crop plants. We used data from 96 records of 62 bee species to determine whether body size predicts foraging distance. We regressed maximum and typical foraging distances on body size and found highly significant and explanatory nonlinear relationships. We used a second data set to: (1) compare observed reports of foraging distance to the distances predicted by our regression equations and (2) assess the biases inherent to the different techniques that have been used to assess foraging distance. The equations we present can be used to predict foraging distances for many bee species, based on a simple measurement of body size. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Dental microwear and diet in Venezuelan primates.

Recent microwear analyses have demonstrated that wear patterns can be correlated with dietary differences. However, much of this work has been based on analyses of museum material where dates and locations of collection are not well known. In view of these difficulties, it would be desirable to compare microwear patterns for different genera collected from the same area at the same time. The opportunity to do this was provided by the collections of the Smithsonian Venezuelan Project (Handley, 1976), in which multiple primate genera were collected from the same humid tropical forest sites within the same month. The monkeys represent a wide range of dietary preferences, and include Saimiri, Cebus, Chiropotes, Ateles, Aotus, Pithecia, and Alouatta. As in previous microwear analyses, epoxy replicas were prepared from dental impressions, as described by Rose (1983) and Teaford and Oyen (1989). Two micrographs were taken of facet 9 on an upper second molar of each specimen. Computations and analyses were the same as described by Teaford and Robinson (1989). Results reaffirm previously documented differences in dental microwear between primates that feed on hard objects versus those that do not--with Pithecia and Alouatta at the extremes of a range of microwear patterns including more subtle differences between species with intermediate diets. The subtle microwear differences are by no means easy to document in museum samples. However, additional results suggest that 1) the width of microscopic scratches may be a poor indicator of dietary differences, 2) large and small pits may be formed differently, and 3) there are very few seasonal differences in dental microwear in the primates at these humid tropical forest sites.     

Prey size-distributions and size-specific foraging success of Ambystoma larvae

Summary We examined how prey size-distributions influence size-specific foraing rate and food gain, i.e., food intake scaled to metabolic demands, in Jefferson's and small-mouth salamander larvae. Ambystoma jeffersonianum larvae sampled on 17 dates from a farm pond whose fauna was dominated by macrozooplankton and chironomid larvae were rarely gape-limited, and total volume of food in the stomach (VS) showed only a slight tendency to increase with larval size. Although 15 of 17 correlation coefficients of VS with larval size were positive, only 1 of 17 correlations were statistically significant, and body size explained only 8% of the overall variation in VS. Correlation coefficients of food gain and body size were positive in 9 cases and negative in 8, but only 3 were statistically significant.In contrast, Ambystoma texanum larvae in 42 samples taken from five sites dominated by macrozooplankton as well as relatively large isopods and amphipods were almost always gape-limited, and VS tended to increase markedly with larval size. 40 of 42 correlation coefficients of VS and larval size were positive, and 19 correlations were statistically significant. Body size in turn explained about 35% of the overall variation in VS. Correlation coefficients of food gain and larval size were positive in 32 of 42 samples, and 9 of 10 significant correlations were positive.When food is limiting and prey selection is not limited by gape, smaller larvae may grow as fast or in some cases faster than larger larvae because they are nearly as effective foragers, but have lower metabolic demands. Larger larvae may in turn grow faster than smaller larvae in environments which support a broad size spectrum of prey, particularly when gape limitations are highly disproportionate among size classes. The growth rate of larvae in one size class relative to another depends primarily on the extent to which increased foraging rate compensates for higher energy demands as body size increases. Size-specific foraging rate may in turn be strongly influenced by the prey size-distribution within a habitat. These relationships suggest that relative size is not always a good a priori predictor of exploitative competitive ability.     

Regulation of foraging trips and incubation routine in male and female wandering albatrosses

The resolution of the conflict between eggcare and foraging was studied in male and female wandering albatrosses. The foraging zone and range, duration of incubation shifts and foraging trips, and associated changes in body mass were studied. Costs during incubation, expressed as the time spent incubating and the proportional loss of body mass, were similar for both sexes. The mass gained at sea was related to the duration of foraging trips, but the relationship was much less significant in males, where foraging ranges, though similar on average to those of females, were very variable. Males foraged in more southerly waters than females, and gained mass more rapidly. Only females appeared to regulate the duration of foraging trips, and this compensated for the mass lost during the incubation fast. Previous breeding experience had no influence on foraging efficiency. Egg desertion because of depletion of body reserves was very rare because birds have a wide safety margin, i.e. the difference between the average body mass when relieved and that at nest desertion. This safety margin enables the birds to compensate for the high variability in the duration of foraging trips, and is probably a reason for the high breeding success of wandering albatrosses. Decisions to return from the sea to the nest or to desert the nest are probably related to the status of body reserves, and have been selected in the large wandering albatross so that both present and future reproductive success are maximised.     

Morphological and behavioral antipredatory adaptations of decapod zoeae

Summary Zoeae of some species of estuarine decapods are retained in the estuary throughout development while others are exported into nearshore coastal waters. The horizontal migrations of decapod zoeae to coastal waters may have evolved to reduce the probability of encountering planktivorous fishes which are most abundant in the estuary. If so, then the morphological vulnerability of zoeae to fish predation should be inversely related to the number of predators occurring where they develop. Six species of estuarine decapod zoeae were offered to Menidia menidia and Fundulus heteroclitus. The behavioral interactions were observed to determine the prey's vulnerability to predation, and the mode of operation and relative effectiveness of their defenses. Feeding trials and behavioral observations both demonstrated that M. menidia 6–16 mm long preferred Uca minax and Callinectes sapidus zoeae, which are exported from the estuary, to Rhithropanopeus harrisii, Sesarma reticulatum and Palaemonetes pugio, which are retained within estuaries. Pinnotheres ostreum zoeae develop in the lower estuary and fish demonstrated an intermediate preference for the zoeae. Menidia menidia 20–40 mm long showed similar preferences for R. harrisii S. reticulatum, P. ostreum and U. minax as did small silversides. Large-mouthed demersal fish, Fundulus heteroclitus 6–10 mm long, also preferred U. minax to R. harrisii, but more readily preyed on zoeae than did M. menidia. The exported species of zoeae have shorter spines and smaller bodies than do retained zoeae, except P. ostreum which is small, spineless and passively sinks when attacked by fish. Other retained species of zoeae also have postcontact behavioral defenses which enhance the effectiveness of their morphological defenses. Zoeae do not evade attacks by fishes, but fishes quickly learned to avoid zoeae, which increases the effectiveness of the zoeae's antipredatory adaptations.     

Systematics and body size: implications for feeding adaptations in New World monkeys.

The relationship between body size and feeding ecology is well established for primates. It is argued that the evolutionary history of modern New World monkeys and, in particular, the path to attainment of current body size is significant in understanding the similarities and differences between dietary strategies and other ecological parameters of similar-sized monkeys. Current interpretations of New World monkey evolutionary relationships are reviewed. Based on a synthesis of available body weights and the assumption that the earliest New World monkeys weighed close to 1 kg, similar to modern Aotus and Callicebus, predicted patterns of body size change in each lineage are given. Restrictions on directions of body size change in primates are discussed, and it is shown that "Stanley's Rule" offers a good explanation for differing body size ranges in New and Old World anthropoids. Predicted ecological correlates to body size drawn from the mammalian literature are offered and tested using data on New World monkeys, which show some concurrence and several interesting departures from predicted patterns. Sexual dimorphism in body weight of New World monkey species is reviewed, based on the new summary of body weight data given.     

Claw morphology, prey size selection and foraging efficiency in generalist and specialist shell-breaking crabs

Claw morphology, and claw-closing forces of four species of intertidal crabs from San Juan Island, Washington were compared and related these findings were related to prey size selection, shell breaking times and total handling times on their snail prey, Littorina sitkana Philippi. Two functional groups of crabs emerged: generalists and specialists on hard-shelled prey. The generalist, Hemigrapsus nudus (Dana), has an omnivorous diet and possesses weak claws with small, fine denticles and mechanical advantage (MA) of the claw's lever system <0.3, while the specialists, Lophopanopeus bellus (Stimpson), Cancer oregonensis (Dana) and C. productus (Randall), consume hard-shelled prey and possess large, powerful claws with broad, blunt molars and MA>0.3. The claws of the generalist, H. nudus, exhibited weaker claw closing forces (5 N) than those of similar sized specialists (>12 N). When crabs of similar weight were offered four size categories of Littorina sitkana, the generalist, Hemigrapsus nudus, exhibited a consistent preference for the smallest size categories, while the three specialists attacked all size classes offered. Hemigrapsus nudus took significantly longer (134 s) than the specialists (30–52 s) to break open a 4 mm L. sitkana. This difference in shell-breaking time between the generalist and the specialists increased with increasing prey size. The rate of successful attacks on increasingly larger L. sitkana decreased with prey size in the generalist (70% on 4 mm, 37% on 6 mm, and 0% on 8 mm snails), but remained high in the specialists (70–100%). Strength limitation of the claws is the best hypothesis to explain the avoidance of large snails by the generalist, H. nudus.     

Experimental increase of flying costs in a pelagic seabird: effects on foraging strategies,nutritional state and chick condition

A central point in life history theory is that parental investment in current reproduction should be balanced by the costs in terms of residual reproductive value. Long-lived seabirds are considered fixed investors, that is, parents fix a specific level of investment in their current reproduction independent to the breeding requirements. We tested this hypothesis analysing the consequences of an experimental increase in flying costs on the foraging ecology, body condition and chick condition in Cory’s shearwaters Calonectris diomedea. We treated 28 pairs by reducing the wing surface in one partner and compared them with 14 control pairs. We monitored mass changes and incubation shifts and tracked 19 foraging trips per group using geolocators. Furthermore, we took blood samples at laying, hatching and chick-rearing to analyse the nutritional condition, haematology, muscle damage and stable isotopes. Eighty-day-old chicks were measured, blood sampled and challenged with PHA immune assay. In addition, we analysed the effects of handicap on the adults at the subsequent breeding season. During incubation, handicapped birds showed a greater foraging effort than control birds, as indicated by greater foraging distances and longer periods of foraging, covering larger areas. Eighty-day-old chicks reared by treated pairs were smaller and lighter and showed a lower immunity than those reared by control pairs. However, oxygen demands, nutritional condition and stable isotopes did not differ between control and handicapped birds. Although handicapped birds had to increase their foraging effort, they maintained physical condition by reducing parental investment and transferred the experimentally increased costs to their partners and the chick. This result supports the fixed investment hypothesis and is consistent with life history theory.     

Reproductive strategies of primates: The influence of body size and diet on litter size

The frequency of multiple births, life history parameters, body size, and diet characteristics were obtained from the literature for 70 primate species. The general pattern within the primate order is to have single infant litters, yet multiple births regularly occur in a number of species in specific phylogenetic groups. Primates which have large litters tend to be small, have short gestation periods and give birth to small infants, which are weaned quickly, and mature rapidly. Species in which multiple births are common also have short interbirth intervals and in the Callitrichidae have males which exhibit paternal care. In addition, they are commonly insectivorous. Although it is difficult to isolate the effects of diet on litter size, independent of body size, analyses suggest that after the influence of body size is statistically removed, as the proportion of insects in the diet increases, animals have larger litters. We suggest that by adopting a mixed diet of insects and fruit primates may be able to ensure access to a seasonally stable food resource that is not greatly restricted by the presence of toxins. This diet would allow a relatively high metabolism and facilitate large litters.     

Effects of prey size and foraging mode on the ontogenetic change in feeding niche ofColostethus stepheni (Anura: Dendrobatidae)

The feeding niche ofColostethus stepheni changes during ontogeny. Small individuals eat small arthropods, principally mites and collembolans, and larger frogs eat bigger prey of other types. The shift in prey types is not a passive effect of selection for bigger prey. There is a strong relationship between electivity for prey types and frog size, independent of electivity for prey size. Four indices of general activity during foraging (number of movements, velocity, total area utilized and time spent moving), which are associated with electivity for prey types in adult frogs and lizards, did not predict the ontogenetic change in the diet ofC. stepheni. Apparently, the behavioral changes that cause the ontogenetic change inC. stepheni are more subtle than shifts in general activity during foraging. Studies of niche partitioning in communities of anurans that do not take into consideration ontogenetic changes in diet and seasonal changes in the size structures of populations present a partial and possibly erroneous picture of the potential interactions among species.     

Group size effects on foraging and vigilance in migratory Tibetan antelope

Large group sizes have been hypothesized to decrease predation risk and increase food competition. We investigated group size effects on vigilance and foraging behaviour during the migratory period in female Tibetan antelope Pantholops hodgsoni, in the Kekexili Nature Reserve of Qinghai Province, China. During June to August, adult female antelope and yearling females gather in large migratory groups and cross the Qinghai-Tibet highway to calving grounds within the Nature Reserve and return to Qumalai county after calving. Large groups of antelope aggregate in the migratory corridor where they compete for limited food resources and attract the attention of mammalian and avian predators and scavengers. We restricted our sampling to groups of less than 30 antelopes and thus limit our inference accordingly. Focal-animal sampling was used to record the behaviour of the free-ranging antelope except for those with lambs. Tibetan antelope spent more time foraging in larger groups but frequency of foraging bouts was not affected by group size. Conversely, the time spent vigilant and frequency of vigilance bouts decreased with increased group size. We suggest that these results are best explained by competition for food and risk of predation.     

Patterns of foraging and range use by three species of neotropical primates

This paper describes the diet and range use patterns of the three species of primates in Santa Rosa National Park, Costa Rica (Ateles geoffroyi, Alouatta palliata, andCebus capucinus) and examines the variation in these variables as they relate to seasonal changes and concomitant changes in food availability. These three primate species were studied over a four-year period for a total of 24 months in the field. Santa Rosa National Park is in an area that experiences a long severe dry season in which little if any rain falls and the majority of the non-riparian trees lose their leaves. However, even though the three species were very flexible in terms of diet and range use, the behavioural variability did not correspond to changes in food availability or season.     

Predator-mediated habitat use: some consequences for species interactions

Synopsis Behavioral responses to predators can have a major impact on a fishes' diet and habitat choice. Studies with the bluegill sunfish, Lepomis macrochirus, demonstrate that bluegills undergo pronounced shifts in diet and habitat use as they grow in response to changes in their vulnerability to predators. Other species of fish exhibit similar habitat shifts with body size, presumably also in response to changing predation risks and/or foraging gains. An important but little appreciated consequence of this type of predator-mediated habitat use is that predation risk, by structuring size and/or age-specific resource use, may also indirectly affect species interactions. This paper discusses some of the ways in which behavioral responses to predators may affect intra- and interspecific competition in fish. Observational and experimental studies with sunfish (Centrarchidae) provide most of the examples. These studies suggest that the nonlethal effects of predators may be as important as the actual killing of prey.     

Competition in foraging flocks of migrating semipalmated sandpipers

I examined the effect of competitor density on foraging success in staging semipalmated sandpipers (Calidris pusilla) foraging on a burrowing amphipod (Corophium volutator) in each of two study years. Little is known about the effect of competitor density when predation attempts disturb prey, causing a temporary decrease in food availability. Controlling for Corophium density and other potentially confounding factors such as temperature, pecking rate and capture rate increased linearly with sandpiper density. Success rate, the ratio of captures to pecks, was not influenced by sandpiper density. The effect of sandpiper density was similar in each of the two study years and was documented early and late in the low tide period. The increase in foraging rate is argued to be a response to increased competition for rapidly depleting prey at the temporal scale of exploitation by a flock. Potential fitness costs associated with higher foraging costs may include decreased ability to distinguish between the profitability of different prey and reduced vigilance against predators.     

Correlation of tooth size and body size in living hominoid primates, with a note on relative brain size in Aegyptopithecus and Proconsul.

Second molar length and body weight are used to test the correlation between tooth size and body size in living Hominoidea. These variates are highly correlated (r= 0.942, p less than 0.001), indicating that tooth size can be used in dentally unspecialized fossil hominoids as one method of predicting the average body weight of species. Based on tooth size, the average body weight of Aegyptopithecus zeuxis is estimated to have been beteen 4.5 and 7.5 kg, which is corroborated by known cranial and postcranial elements. Using Radinsky's estimates of brain size, the encephalization quotient (EQ) for Aegyptopithecus was between 0.65 and 1.04. A similar analysis for Proconsul africanus yields a body weight between 16 and 34 kg, and an EQ between 1.19 and 1.96.     

Tubes and foraging behavior in larval Chironomidae: implications for predator avoidance

Summary In laboratory experiments, I studied differential susceptibility of four co-occurring species of chironomids to a predatory damselfly. The chironomids differed in foraging behavior and could be ranked according to the amount of time they spent outside of their tubes. In choice experiments, the predator consistently selected the prey which spent more time out of the tube, and time out of tube was a significant predictor of the predation rate coefficient. Electivity indices, calculated from field samples and diet analyses of the predator, supported the laboratory results. The data suggest that exposure to predators in a heterogeneous prey community is largely determined by tubedwelling behavior.