Spatial position and feeding success in ring-tailed coatis

The location of an animal within a social group has important effects on feeding success. When animals consume quickly eaten food items, individuals located at the front edge of a group typically have greater foraging success. When groups feed at large clumped resources, dominant individuals can often monopolize the resource, leading to higher feeding success in the center of the group. In order to test these predictions, behavioral data relating foraging success to within-group spatial position were recorded from two habituated groups of ring-tailed coatis (Nasua nasua) in Iguazu, Argentina. Foraging success did not fit expected patterns. When feeding on small ground litter invertebrates, coatis had the same foraging success at all spatial positions. This pattern likely resulted from an abundance of invertebrates in the ground litter. When feeding on fruit, individuals in the front of the group had greater feeding success, which was driven by the relatively quick depletion of fruit trees. Dominant juveniles were often located in the front of the group which led to increased access to food. This resulted in higher feeding success on fruits but simultaneously increased their risk of predation. Although groups typically became more elongated and traveled faster when feeding on fruit, it did not appear that the coatis were drastically changing their spacing strategies when switching between the two food types. Paradoxically, spatial position preferences during invertebrate foraging appeared to be driven by fruit trees. Because fruit trees were encountered so frequently, juveniles ranging at the front edge of the group during invertebrate foraging were the first to arrive at fruit trees and thus had higher foraging success. This study demonstrates the importance of how food patch size and depletion rate affect the spatial preferences of individuals.     

Do fledglings choose wisely? An experimental investigation into social foraging behaviour

Many cooperative bird species have an extended period of post-fledging care. Despite the fact that this period of care can last up to several months, it remains a relatively understudied stage of chick development. This period, when young are actively begging but highly mobile, provides an opportunity for young to maximise the amount of care they receive by selectively choosing particular adults to beg from. In pied babblers Turdoides bicolor (a cooperatively breeding passerine), fledglings closely follow foraging adults and beg for food regularly (a behavioural interaction termed social foraging). Using a combination of natural observations and experimental manipulations, we found that fledgling pied babblers preferentially socially forage with adult care-givers who have high foraging success, since this results in young receiving more food. By supplementally feeding adults to artificially increase their foraging success, we increased the proportion of time that fledglings chose to socially forage with them, confirming that fledglings are selectively choosing dyadic interactions with the best adult foragers. These results indicate that pied babbler fledglings are sensitive to and can respond to short-term changes in adult foraging success, enabling them to maximize their nutritional intake, a behavioural adjustment that has long-term benefits in this system.     

The association of female kin in the arctic ground squirrel Spermophilus parryii

Summary The association of kin in arctic ground squirrels (Spermophilus parryii) was studied near Haines Junction, Yukon Territory, during the summers of 1977, 1978, and 1979. Males dispersed in this population, thus only females were likely to live near relatives.Close female kin (sisters, mothers/daughters) had greater overlap of home ranges and interacted more amicably and less agonistically than did less closely related females. Closely related females clumped their young at emergence, whereas more distantly related females did not; there was little indication of clumping of pre-emerged young. The overlap of home ranges of distant relatives (known genetic relatives that had not associated in a natal burrow) was intermediate to that of close relatives and non-relatives. The types of interactions between distant relatives were more similar to those between non-relatives than between close relatives.I conclude that female arctic ground squirrels exhibit nepotism. Females may benefit from associations with relatives by sharing watching for predators once juveniles become conspicuous. Male arctic ground squirrels commit infanticide and several females may be more effective at protecting their young from infanticidal males than females living alone. I suggest that clumping of young by close relatives may provide a mechanism allowing distantly related females to identify each other.Address for offprint requests     

Individual variation in winter foraging of black-capped chickadees

Summary Wintering black-capped chickadees (Paridae: Parus atricapillus) in northwestern Massachusetts showed a high degree of individual variation in foraging behavior. After accounting for the effects of different habitats and weather conditions, individual differences comprised 6–17% of the total observed variation in four measures of foraging location and rate of feeding. Differences between age and sex groups were not significant and explained comparatively little variation (0.0–1.4%). The chickadees did not fall into a few distinct behavioral categories but instead showed continuous variation on all measures of foraging behavior. It appeared that some variation among individuals was a consequence of behavioral convergence within social groups, since birds that were observed together were more similar in their foraging than expected by chance, after taking habitat differences into account. Our results therefore do not support the interpretation that individual variation in feeding behavior serves to reduce exploitation competition within social groups.     

Feeding enhancement by social attraction in the Sandwich Tern

Summary A controversy exists over the relative roles of feeding enhancement and predation in shaping the flocking behavior of birds. We examined social attraction among foraging Sandwich Terns, Sterna sandvicensis, in the Gulf of Mexico. The prediction that foraging terns would coalesce at a spot where another bird dove was supported. Approach took place within seconds, occurred over a distance of at least 500 m, and lasted about 10 to 20 s after the food source dissipated. Although social attraction appears to play a major role in the feeding of the terns, this need not be construed as evidence against a similarly important role for predation in shaping social foraging behavior.     

Why do female Belding's ground squirrels disperse away from food resources?

We examined the effects of food provisioning on the natal dispersal behavior of Belding's ground squirrels (Spermophilus beldingi). We provided extra food to adult and yearling females in their maternal territories during pregnancy and lactation, and to offspring of these females in their natal areas for 6 weeks after weaning. We used unprovisioned young of unprovisioned mothers as controls. Provisioning influenced the probability of dispersal from the natal area by female but not male S. beldingi. All surviving male S.␣beldingi dispersed by 55 weeks of age, regardless of whether they and their mothers received extra food. By contrast, we observed a significant trend, beginning 3 weeks after weaning and continuing through the yearling year, for a greater proportion of provisioned than control female S. beldingi to emigrate from the natal area. Competition for food did not appear to influence natal dispersal of females. However, overall population density, density of females weaning litters, and rates of aggression and vigilance among these females, were higher in provisioned than control areas, suggesting that competition for non-food resources was unusually intense in provisioned areas. We propose that juvenile female, but not juvenile male, S. beldingi may emigrate from the natal site to increase access to areas with low densities of conspecifics. Together with findings of earlier workers, our results suggest that spatial and temporal distributions of environmental resources are important influences on the dispersal behavior of female ground squirrels. Received: 28 February 1996 / Accepted after revision: 16 October 1996     

Subhabitat selection by foraging threespine stickleback (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>): previous experience and social conformity

Any mechanism that allows animals to increase their foraging efficiency is likely to be selected for, including the ability to learn to recognise and subsequently discriminate between habitat types based on their profitability. In a series of laboratory studies, we manipulated prey densities across two different experimental subhabitats and demonstrated that threespine stickleback (Gasterosteus aculeatus) can develop foraging preferences for subhabitats that have previously yielded prey. Fish were not recalling the spatial location of prey patches; rather, they were discriminating between subhabitats based on foraging experience there and allocating foraging effort accordingly. Foraging preferences took around 14 days to develop, and once established, they persisted independently of experimental prey density, suggesting that fish were using experience rather than real-time sampling to select foraging grounds. When we presented focal fish with social information cues, we found that they preferentially used local enhancement and current public information cues when they conflicted with previous experience, but that they did not use prior public information. This suggests that in the presence of conspecifics, individuals prioritise social conformity over the use of private information. We discuss our results in the context of optimal foraging and the trade-offs associated with balancing conflicting private and social information.     

Variation in foraging effort by lactating Antarctic fur seals: response to simulated increased foraging costs

Seasonally breeding predators, which are limited in the time available for provisioning young at a central location, and by the fasting abilities of the young, are likely to maximize energy delivery to the young by maximizing the rate of energy delivery averaged over the whole period of investment. Reduction in food availability or increased foraging costs will alter the optimal behavior of individuals. This study examined the behavioral adaptations of a diving predator, the Antarctic fur seal, to increased foraging costs during lactation. One group of mothers (n=5, treatment) was fitted with additional drag to increase the cost of transport in comparison with a control group (n=8). At the scales of the individual dives, the treatment group made more shorter, shallower (< 30 m) dives. Compensation for slower swimming speeds was achieved by diving at a steeper angle. Overall, diving behavior conformed to several specific theoretical predictions but there were also departures from theory, particularly concerning swimming speed during diving. Diving behavior appears to be adjusted to maximize the proportion of time spent at the bottom of dives. At the scale of diving bouts, no difference was observed between the treatment and control groups in terms of the frequency and duration of bouts and there was also no difference between the two groups in terms of the proportion of time spent diving. At the scale of complete foraging cycles, time taken to return to the pup was significantly longer in the treatment group but there was no difference in the rate of delivery of energy (measured from pup growth rate) to the pups in each group. Since mothers in the treatment group did not use significantly more body reserves, we conclude that behavioral adjustments at the scale of individual dives allowed mothers in the treatment group to compensate for the additional foraging costs. Pup growth rate appears to be less sensitive to the foraging conditions experienced by mothers than foraging trip duration. Received: 14 June 1996 / Accepted after revision: 16 November 1996     

Nutritional constraints on mountain baboons (Papio ursinus): Implications for baboon socioecology

Summary Populations of baboon (Papio sp.) at geographic and climatic extremes for the genus show a tendency to one-male organization, whereas most baboons live in multimale social groups; this effect has been attributed largely to limitation of food supply, but baboons' complex diet has hindered proper nutritional analyses. To test these optimal-diet explanations of social variation, we quantified intake and used phytochemical analysis of foods to compare the nutrition, during seasonal changes, of two groups of mountain baboons (P. ursinus) living at different altitudes of a continuous grassland habitat. The majority of plant foods were eaten uniquely by one or other group, though their altitudinal separation was only 400 m, and the time budget of feeding choices varied with age-sex class as well as season. Converting to a common currency of nutrients reveals that baboons gained the same yield from a unit time spent foraging (whether this is measured in edible dry weight, or simply protein) in both groups, despite their differing mean altitude, whereas seasonal variation was large and statistically significant. Increased feeding time at the winter bottleneck made no effective compensation for the poorer food yields: in late winter there was a minimum for daily nutrient gain at both altitudes. Apparently this population is already at an extreme for the time animals devote to foraging in winter, when they rely on inconspicuous and slow-to-harvest swollen shoot bases and underground plant storage organs. Since an individual's nutrient yield does not vary with altitude, we conclude that socioecological parameters are effectively optimized for feeding. Since contest competition is absent, this adjustment of foraging efficiency is largely through the effect of differential density on scramble competition. Differences in social structure are considered to be a secondray consequence of optimal foraging, mediated through altitudinal variation in either population density or in day range limits.     

Individual energetic state can prevail over social regulation of foraging in honeybees

The energetic state of an individual is a fundamental driver of its behavior. However, an individual in a eusocial group such as the honeybees is subject to the influence of both the individual and the colony energetic states. As these two states are normally coupled, it has led to the predominant view that behaviors, such as foraging, are dictated by the colony state acting through social regulatory mechanisms. Uncoupling the energetic state of an individual honeybee from its colony by feeding it with a non-nutritious sugar, we show that energetically stressed bees in a colony with full food stores do not consume this food to meet their energetic shortfall but instead compensate by first reducing their activity level and then by increasing their foraging rate. This suggests that foraging in eusocial groups is still partly under the regulatory control of the energetic state of the individual and supports the notion that regulatory mechanisms in solitary insects have been co-opted to drive altruistic behavior in eusocial insects. The observation that energetically stressed bees also experience higher mortality during foraging also suggests that energetic stress mediated by a variety of factors can be a common mechanism that underlies the recent observation of bees disappearing from their colonies. We also discuss how nutritional imbalance in a social insect individual can alter its behavior to influence colony life history.     

The pit-trapping foraging strategy of the ant lion,Myrmeleon immaculatus DeGeer (Neuroptera: Myrmeleontidae)

Summary Larvae of Myrmeleon immaculatus in large pits captured both large and samll prey, while larvae in small pits captured only the small prey. Larvae in small pits did not respond to large ants, although they always responded by sand-flinging to small ants. Larvae in medium-sized pits often captured large ants only after prolonged and vigorous sand-flipping. Larvae in large pits usually captured large ants with relatively little sand-flipping. Pit enlargement and pit relocation in the laboratory were not significantly correlated with reduction of rations in the first 3 weeks after a pit was built. However, after a month without food, larvae on the average moved once every 10 days, built successively smaller pits, and moved longer distances before building a new pit. In the field pits were dug primarily in response to microclimatological factors and possibly edge-effects. The presence or absence of suitable prey at a site, per se, had no effect on whether or not a larva would dig a pit there. We conclude that these sit-and-wait predators have a relatively large repertoire of behavior that enhances their foraging success, and we contrast it with previously made optimal foraging models relating to pit locations, pit relocations, pit size and ant lion responses.     

Division of labor between undertaker specialists and other middle-aged workers in honey bee colonies

A primary determinant of colony organization in temporally polyethic insect societies is inter-individual variation in behavior that is independent of worker age. We examined behavioral repertoires, behavioral correlates of adult development, and spatial distributions within the hive to explore the mechanisms that produce behavioral variation among middle-age honey bees (Apis mellifera). Individually labeled undertakers, guards, food storers, and wax workers exhibited a broad range of task-related behavior, but bees tagged as undertakers were more likely to subsequently remove a corpse from the hive and handle a corpse compared to other middle-aged bees. The activity level of undertakers was similar to other task groups, suggesting that undertaking specialists were neither hyper-active “elites” nor quiescent “reserves” that become active only when a dead bee stimulus is present. Undertakers also were more likely to remove debris and to remain in the lower region of the hive or near the entrance, even when not engaged in corpse removal; both preferences may promote colony efficiency by reducing inter-task travel times. Guards and undertakers were less likely to perform behavior normally associated with young bees compared to food storers and wax workers. Undertakers and guards also initiated foraging at earlier ages than the other task groups. These results suggest that undertakers and guards may be slightly developmentally advanced compared to food storers and wax workers. There also was evidence for lifetime differences in behavioral preferences which could not be explained by differences in adult development. Bees tagged as undertakers were more likely to subsequently remove a dead bee during their entire pre-foraging career compared to other task groups or members of their general age cohort. Differences in both the rate of adult development and individual behavioral preferences, both of which may be subject to genetic and environmental influences, are important determinants of inter-individual variation among honey bees of middle age. Received: 5 February 1997 / Accepted after revision: 27 May 1997     

Collective foraging decision in a gregarious insect

Group foraging by eusocial insects implies sophisticated recruitment processes that often result in collective decisions to exploit the most profitable sources. These advanced levels of cooperation, however, remain limited to a small range of species, and we still know little about the mechanisms underlying group foraging behaviours in the great mass of animals exhibiting lower levels of social complexity. In this paper, we report, for the first time in a gregarious insect, the cockroach Blattella germanica (L.), a collective foraging decision whereby the selection of food sources is reached without requiring active recruitment. Groups of cockroaches given a binary choice between identical food sources exhibited exploitation asymmetries whose amplitude increases with group size. By coupling behavioural observations to computer simulations, we demonstrate that selection of food sources relies uniquely on a retention effect of feeding individuals on newcomers without comparison between available opportunities. This self-organised pattern presents similarities with the foraging dynamics of eusocial species, thus stressing the generic dimension of collective decision-making mechanisms based on social amplification rules despite fundamental differences in recruitment processes. We hypothesise that such parsimony could apply to a wide range of species and help understand the emergence of collective behaviours in simple social systems.     

Reward rate and forager activation in honeybees: recruiting mechanisms and temporal distribution of arrivals

We analyzed the foraging and recruitment activity of single foragers ( Apis mellifera), exploiting low reward rates of sucrose solution. Single employed foragers (test bees) were allowed to collect 2.0 m sucrose solution delivered by a rate-feeder located at 160 m from the hive for 2 h. Flow rates varied between 1.4 and 5.5 µl/min. The individual behavior of the test bees was registered both at the hive and the food source, and the social output was calculated as the number of incoming bees arriving at the feeder per hour (henceforth: arrival rate). Incoming bees were captured once they landed at the feeder and assigned to one of three categories according to their foraging experience and hive interactions with the test bee: inspector, reactivated, or inexperienced bees. Both the waggle-runs performed per hour of foraging by test bees and the social output attained, increased with the reward rate. Also the number of hive-stays and the trophallactic-offering contacts performed by test bees were positively correlated with the arrival rate. For the highest reward rates, the duration of Nasonov-gland exposure at the feeding place was higher, and the arrival of most of the incoming bees occurred shortly after the test bee landed at the feeding platform. Thus, in addition to hive-interactions, landing of incoming bees at the food source is promoted by olfactory and/or visual information provided by the test bees. The proportions of inspector, reactivated, and inexperienced bees changed depending on the reward rate offered. Therefore, not only the occurrence and intensity of the recruitment-related behaviors performed by the test bees, but also the stimulation required by each category of incoming bees, determined the social output observed.     

Fish feeding on mobile benthic invertebrates: influence of spatial variability in habitat associations

Patterns of habitat association and foraging were examined for a group of tropical goatfishes (family Mullidae) that feed on mobile benthic invertebrates at Lizard Island (Great Barrier Reef). All goatfish possess barbels that disturb the substratum during feeding. Foraging methods were examined for the six most common species and used in conjunction with data on habitat associations to estimate the distribution and potential impact on the benthic invertebrate assemblage of foraging-related disturbance. Particular species exhibited broad habitat associations which differed little over two surveys (January 1989, January 1990). All species showed different preferences for the substrata they foraged. Preferences for substrata exhibited by the most common reef-associated species, Parupeneus multifasciatus, differed among locations separated by 1 km, between sites 150 m apart, and between depths (shallow and deep). Habitat preferences changed with ontogeny. Based on their habitat associations and foraging preferences, species were divided into habitat generalists and specialists. Specialists associated primarily with soft sediments. Habitat generalists, such as P. multifasciatus and P. cyclostomus, are likely to have an impact on their mobile invertebrate prey that is localised, diffuse and transitory, making any experimental analysis difficult and expensive. Habitat specialists form a guild of fishes with complementary feeding modes that efficiently exploit soft sediments and are more amenable to experimental manipulation. Experiments designed to detect the impact of foraging by these fishes must be repeated at different locations and times and must account for depth differences in foraging pressure.     

Behavioral responses of territorial red squirrels to natural and experimental variation in population density

The relative scarcity of studies at the intersection of behavioral and population ecology is surprising given the presumed importance of behavior in density-dependent population regulation. Here we tested whether North American red squirrels (Tamiasciurus hudsonicus) adjust their behavior in response to local population density and whether they use rates of territorial vocalizations in their local neighborhood to assess density. We examined these relationships using 18?years of live trapping and 20?years of behavioral data that were collected across natural variation in local population density. To disentangle the effects of population density on behavior from those due to changes in per capita food abundance or changes in the frequency of antagonistic interactions with neighbors, we also experimentally manipulated population density with long-term food supplementation as well as perceived population density with long-term playbacks of territorial vocalizations. The frequency with which squirrels emitted territorial vocalizations was positively associated with local population density. In contrast, antagonistic physical interactions observed between squirrels and territorial intrusions were rare and the frequency of intrusions was weakly and negatively, not positively, associated with population density. Squirrels experiencing naturally and experimentally high density conditions spent less time in the nest and feeding but more time being vigilant. Similar density-dependent changes in behavior were observed in response to our manipulations of perceived population density, indicating that vocalization rates and not physical interactions or food abundance were the mechanism by which squirrels assessed and responded behaviorally to changes in local density.     

Interspecific competition in tree squirrels: do introduced grey squirrels (<Emphasis Type="Italic">Sciurus carolinensis</Emphasis>) deplete tree seeds hoarded by red squirrels (<Emphasis Type="Italic">S. vulgaris</Emphasis>)?

Red squirrels (Sciurus vulgaris) and introduced eastern grey squirrels (S. carolinensis) scatterhoard seeds of broadleaf trees. Scatterhoarded seeds are an essential resource in spring and their consumption increases red-squirrel fitness. We examined whether grey squirrels partly deplete the high-energy food resources cached by red squirrels, reducing their consumption, in two ways: (1) at the population level, comparing energy intake of feeding on cached seeds between a study site with red and grey squirrels and one with only red squirrels present; and (2) at the individual level, in the study site where species co-exist, relating hoard recovery of red squirrels to the amount of core-area overlap with grey squirrels. There were no significant site differences in the mean daily energy intake of red squirrels feeding on seeds recovered from caches. However, in the red-grey site, during spring, red squirrels that had a high percentage of their home-range core area overlapped by grey squirrels had a lower daily energy intake than low-overlap red squirrels. Body mass of red squirrels in spring was negatively correlated with the percentage of interspecific core-area overlap, but not with core-area overlap with other red squirrels. Our data suggest that interspecific competition for scatterhoarded seeds, with grey squirrels pilfering red squirrels' food caches, caused a reduced energy intake in red squirrels with a high degree of interspecific core-area overlap, and reduced body mass in spring. Therefore, cache pilfering is likely to reduce reproductive output in red squirrels, and thus play a role in the replacement of red by grey squirrels. Electronic Publication     

Transmission of food preferences in spiny mice (Acomys cahirinus) via nose-mouth interaction between mothers and weanlings

Summary Two laboratory experiments assessed the role of nose-to-head and nose-mouth investigation in the transmission of food preferences from Acomys cahirinus mothers to their offspring. Pups spent more time nosing and licking the nose-mouth area of mothers who ate a novel food (fresh potato or carrot) than of mothers who ate no novel food. A greater amount of the less-preferred novel food (potato) was subsequently eaten by pups whose mothers had eaten potato than pups whose mothers had eaten either carrot or familiar laboratory chow only. Mothers maintained a stereotypic posture during nose-mouth investigation, holding still with mouth slightly open, providing both a source of specific food cues and a context in which pups could become familiar with them. These data suggest an active role by Acomys mothers in the development of offspring food preferences.     

The effect of reproductive condition on the foraging behavior of female hoary bats,Lasiurus cinereus

Summary Female mammals experience larg changes in time and energy budgets associated with reproduction and these may influence the foraging strategies of individuals. I studied the changes in foraging behavior associated with reproduction in female hoary bats, Lasiurus cinereus. As lactation progressed, individuals departed to forage earlier in the evening and spent more time foraging per night and less time roosting with their young. Foraging time increased by at least 73% between early lactation and fledging and then declined as the young became independent. Females with two young foraged for longer than did those with one and females with pre- and postfledging young foraged in different habitats. The changes in foraging time suggest that foraging activity of female L. cinereus is constrained and individuals act as time minimizers, adjusting their foraging behavior to meet current energy demand. Predation risk is unlikely to constrain the behavior of these bats. However, maximizing energy intake throughout lactation may not be the optimal strategy because storing excess energy increases flight cost and may reduce foraging efficiency. The need to keep newborn young warm may also influence foraging time. Such constraints, causing changes in foraging activity, may alter the availability of habitats and prey and must be considered when modelling foraging strategies. In addition, changes in flight time may significantly alter the energy budgets of bats in different stages of reproduction.     

Biotic resistance via granivory: establishment by invasive, naturalized, and native asters reflects generalist preference

Escape from specialist natural enemies is frequently invoked to explain exotic plant invasions, but little attention has been paid to how generalist consumers in the recipient range may influence invasion. We examined how seed preferences of the widespread generalist granivore Peromyscus maniculatus related to recruitment of the strongly invasive exotic Centaurea stoebe and several weakly invasive exotics and natives by conducting laboratory feeding trials and seed addition experiments in the field. Laboratory feeding trials showed that P. maniculatus avoided consuming seeds of C. stoebe relative to the 12 other species tested, even when seeds of alternative species were 53-94% smaller than those of C. stoebe. Seed addition experiments conducted in and out of rodent exclosures revealed that weakly invasive exotics experienced relatively greater release from seed predation than C. stoebe, although this was not the case for natives. Seed mass explained 81% of the variation in recruitment associated with rodent exclusion for natives and weak invaders, with larger-seeded species benefiting most from protection from granivores. However, recruitment of C. stoebe was unaffected by rodent exclusion, even though the regression model predicted seeds of correspondingly large mass should experience substantial predation. These combined laboratory and field results suggest that generalist granivores can be an important biological filter in plant communities and that species-specific seed attributes that determine seed predation may help to explain variation in native plant recruitment and the success of exotic species invasions.