Elevation-related differences in novel environment exploration and social dominance in food-caching mountain chickadees

Harsh and unpredictable environments have been assumed to favor the evolution of better learning abilities in animals. At the same time, individual variation in learning abilities might be associated with variation in other correlated traits potentially forming a behavioral syndrome. We have previously reported significant elevation-related differences in spatial memory and the hippocampus in food-caching mountain chickadees. Here, we tested for elevation-related differences in novel environment exploration, neophobia, and social dominance—behavioral traits previously thought to correlate with individual variation in cognition, using different birds from the same elevations. Compared to low-elevation birds, high-elevation chickadees were slower at novel environment exploration, but there were no detectable differences in neophobia. High-elevation chickadees were also socially subordinate to low-elevation chickadees in pairwise interactions. Considering previously reported elevation-related differences in cognition and the brain, our results suggest, however indirectly, that elevation-related variation in spatial memory might be associated with differences in novel environment exploration and in ability to obtain a high social rank in winter social groups. Whether these behavioral traits represent a behavioral syndrome or whether climate might affect these traits independently, our results suggest that multiple differences between elevations might assist with elevation-related separation. High-elevation chickadees would likely experience higher mortality if they move to lower elevation due to their low social dominance status and low-elevation chickadees might experience higher mortality if they move to higher elevation due to reduced memory ability and lack of behavioral adaptations to colder climate.     

Can environmental heterogeneity explain individual foraging variation in wild bottlenose dolphins (<Emphasis Type="Italic">Tursiops</Emphasis> sp.)?

Because behavioral variation within and among populations may result from ecological, social, genetic and phenotypic differences, identifying the mechanism(s) responsible is challenging. Observational studies typically examine social learning by excluding ecological and genetic factors, but this approach is insufficient for many complex behaviors associated with substantial environmental variation. Indian Ocean bottlenose dolphins (Tursiops sp.) in Shark Bay, Western Australia show individual differences in foraging tactics, including possible tool use with marine sponges and social learning may be responsible for this diversity. However, the contributions of ecological factors to the development of these foraging tactics were not previously investigated. Here, we determined the relationship between ecological variables and foraging tactics and assessed whether differences in habitat use could explain individual differences in foraging tactics. We monitored 14 survey zones to identify how foraging tactics were spatially distributed and matched behavioral data to the ecological variables within each zone. Three of four foraging tactics were significantly correlated with ecological characteristics such as seagrass biomass, water depth, presence of marine sponges and season. Further, individual differences in habitat use were associated with some tactics. However, several tactics overlapped spatially and previous findings suggest demographic and social factors also contribute to the individual variation in this population. This study illustrates the importance of environmental heterogeneity in shaping foraging diversity and shows that investigating social learning by ruling out alternative mechanisms may often be too simplistic, highlighting the need for methods incorporating the relative contributions of multiple factors.     

Foraging in a variable environment: weather patterns and the behavioral ecology of baboons

We investigated the long-standing premise in behavioral ecology that the environment affects behavior and demography. We did this by evaluating the extent to which year-to-year variability in the behavioral ecology of a nonhuman primate population could be modeled from meteorological patterns. Data on activity profiles and home range use for baboons (Papio cynocephalus) in Amboseli, Kenya, were obtained over a 10-year period for three social groups: two completely wild-foraging ones, and a third that supplemented its diet with refuse from a nearby tourist lodge. The relationships across years among activity budgeting, travel distance, group size, and measures of temperature and rainfall patterns differed among the social groups. Although meteorological variation generally correlated with behavioral variation in the completely wild-foraging groups, different weather variables and direction of relationships resulted for each group. In addition, different relationships among variables were found before and after home-range shifts. The food-enhanced group spent half as much time foraging as did the other groups and therefore could be used to evaluate the relative extent to which foraging time was a limiting factor for resting and social time. Under their relaxed ecological conditions, the food-enhanced animals increased resting time much more than social time. These findings, combined with supplementary information on the population, lead us to suggest that baboons use a suite of interrelated responses to ecological variability that includes not only changes in activity budgets, but also home-range shifts, changes in the length of the active period, and changes in group size through fissions. Moreover, our results imply that group differences as well as interpopulational and interspecific differences in behavioral ecology provide significant sources of variability. Therefore, social groups rather than populations may be the appropriate unit of analysis for understanding the behavioral ecology of baboons and other highly social primates. The different patterns we observed among groups may have fitness consequences for the individuals in those groups and thereby affect population structure over time. Received: 18 February 1995/Accepted after revision: 6 January 1996     

Effects of body mass,age, dominance and parasite load on foraging time of bighorn rams,<Emphasis Type="Italic"> Ovis canadensis</Emphasis>

In sexually dimorphic ungulates, males generally spend less time foraging than females, possibly because of difference in body mass or because of the energetic requirements of lactation. The relationship between body size and foraging time has received little attention at the intra-specific level, because few studies have documented activity budgets for individuals of known size. Bighorn rams are a good model to explore how body mass affects foraging time, because they range in mass from 55 to 140 kg. We examined how the foraging time of bighorn rams varied according to individual characteristics. We observed rams in a marked population and constructed time budgets during the 3 months preceding the rut. We determined ram social rank based on agonistic encounters and collected fecal samples to count lungworm larvae. Time spent foraging was negatively correlated with body mass. After accounting for age differences, larger rams spent less time foraging and more time lying than smaller rams. Among rams aged 6–12 years, dominants spent less time feeding than subordinates, while fecal output of lungworm larvae was negatively correlated with foraging time for rams of all ages. Body mass accounts for much of the individual variation in foraging time, suggesting that sexual dimorphism is important in explaining differences in feeding time between males and females.Communicated by P. Heeb     

Time budgets and foraging in a Malagasy primate: do sex differences reflect reproductive condition and female dominance?

Female mammals commonly employ behavioral tactics of modulating activity levels and foraging behavior to counter the energetic burden of reproduction; these behavioral changes are reflected as intersexual differences. Traditional views of Malagasy primates posit that high reproductive costs select for female dominance which guarantees to energetically stressed females priority of resource access. I tested predictions regarding reproductive influences on sex differences in time budgets and foraging behavior using two groups of Milne-Edwards' sifaka (Propithecus diadema edwardsi) in southeastern Madagascar. Compared to males, females increased neither feeding nor resting time during gestation or lactation. Sex differences were essentially absent in all foraging time variables examined (time, duration, rate). In contrast, dietary composition diverged between the sexes in some months. The possibility that females selected particular food items to boost nutrient and energetic intake to meet increased requirements during reproduction must be further clarified with nutritional analyses. Sex differences in plant part choices coincided with lactation in one of the two study groups. Thus, the timing of sex differences in feeding patterns of P. d. edwardsi only partially supports the prediction that sex differences are most pronounced during the period of greatest female energetic demand. A comparative review indicated no tight association between female dominance and sex differences in foraging among Malagasy primates. Traditional female dominance theory falls short of explaining the observed patterns. The results of my study coupled with recent evidence suggest that non-behavioral tactics involving energy conservation and storage require further attention as mechanisms by which female lemurs cope with reproductive costs. Received: 12 June 1998 / Accepted after revision: 10 October 1998     

Individual differences and within-flock convergence in chickadee calls

Summary A detailed sound analysis of the Chick-adee call of the black-capped chickadee (Parus atricapillus) was performed in order to determine a basis for individual recognition and for imitation within winter flocks. During the winter of 1978–1979 members of five free-living black-capped chickadee flocks were uniquely marked for individual identification, and their calls were recorded in the field. Nested analysis of variance of temporal call parameters measured from sonagrams and spectral parameters from frequency spectra showed that there were significant differences between individuals within flocks for every parameter measured. There were significant differences between flocks in the frequency ranges 800–2,200 Hz and 4,000–5,300 Hz, in the spectral parameters bandwidth and maximum frequency, and in the duration of the Dee syllable and total call duration.Members of four of the five flocks were captured in December 1978 and held in aviaries segregated by original flock membership. In January 1979 the memberships of the three aviaries were rearranged to form experimental flocks. After one month, there were significant differences among the experimental flocks in the duration of the Dee syllable and total call duration. Convergence, as indicated by a significant decrease in variance among members (F-test), occurred in the experimental flock in Aviary 1 and was concentrated in the frequency ranges 1,300–1,800 and 6,200–6,900 Hz. The members of this experimental flock differed from each other in the number of 100 Hz frequency intervals within which each changed its own call.The Chick-a-dee call contains sufficient information that it can potentially be used by black-capped chickadees for individual recognition. In addition, both field and aviary data suggest that flock members converge in some call characteristics. Possible explanations of the social significance of vocal convergence in chickadee flocks are discussed.     

Division of labor in a dynamic environment: response by honeybees (Apis mellifera) to graded changes in colony pollen stores

 A fundamental requirement of task regulation in social groups is that it must allow colony flexibility. We tested assumptions of three task regulation models for how honeybee colonies respond to graded changes in need for a specific task, pollen foraging. We gradually changed colony pollen stores and measured behavioral and genotypic changes in the foraging population. Colonies did not respond in a graded manner, but in six of seven cases showed a stepwise change in foraging activity as pollen storage levels moved beyond a set point. Changes in colony performance resulted from changes in recruitment of new foragers to pollen collection, rather than from changes in individual foraging effort. Where we were able to track genotypic variation, increases in pollen foraging were accompanied by a corresponding increase in the genotypic diversity of pollen foragers. Our data support previous findings that genotypic variation plays an important role in task regulation. However, the stepwise change in colony behavior suggests that colony foraging flexibility is best explained by an integrated model incorporating genotypic variation in task choice, but in which colony response is amplified by social interactions. Received: 17 October 1998 / Received in revised form: 11 March 1999 / Accepted: 12 March 1999     

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     

Behavioral correlations across activity, mating, exploration, aggression, and antipredator contexts in the European house cricket, Acheta domesticus

Recently, there has been increasing interest in behavioral syndrome research across a range of taxa. Behavioral syndromes are suites of correlated behaviors that are expressed either within a given behavioral context (e.g., mating) or between different contexts (e.g., foraging and mating). Syndrome research holds profound implications for animal behavior as it promotes a holistic view in which seemingly autonomous behaviors may not evolve independently, but as a “suite” or “package.” We tested whether laboratory-reared male and female European house crickets, Acheta domesticus, exhibited behavioral syndromes by quantifying individual differences in activity, exploration, mate attraction, aggressiveness, and antipredator behavior. To our knowledge, our study is the first to consider such a breadth of behavioral traits in one organism using the syndrome framework. We found positive correlations across mating, exploratory, and antipredatory contexts, but not aggression and general activity. These behavioral differences were not correlated with body size or condition, although age explained some of the variation in motivation to mate. We suggest that these across-context correlations represent a boldness syndrome as individual risk-taking and exploration was central to across-context mating and antipredation correlations in both sexes.     

Predator stimuli and calling behavior of Carolina chickadees (Poecile carolinensis), tufted titmice (Baeolophus bicolor), and white-breasted nuthatches (Sitta carolinensis)

Evidence from different chickadee species (Poecile genus) indicates that birds can modify the note composition of their “chick-a-dee” calls in the presence of predator stimuli. Here, we tested the effects of predator models and the distance of those models on calls of three species foraging together at feeding stations: Carolina chickadees (Poecile carolinensis) and tufted titmice (Baeolophus bicolor), both members of the Paridae family, and white-breasted nuthatches (Sitta carolinensis), a member of the Sittidae family. Model and distance affected seed-taking rates in all three species. “Chick-a-dee” calling rates were higher in the predator context for both chickadees and titmice, but we detected no predator context effects on “quank” call rates for nuthatches. Predator and distance contexts affected acoustic parameters of notes of the “chick-a-dee” calls of chickadees and titmice; no such effects were detected for nuthatch “quank” calls. These results suggest species differences in encoding of information in the primary social calls of these three species that commonly occur in multi-species flocks. Chickadees and titmice are “nuclear” species and nuthatches are “satellite” species, and these different roles might be related to the differences in vocal signaling that we detected.     

Group size and foraging efficiency in yellow baboons

Summary I studied the foraging behaviour of adults in three different-sized groups of yellow baboons (Papio cynocephalus) at Amboseli National Park in Kenya to assess the relationship between group size and foraging efficiency in this species. Study groups ranged in size from 8 to 44 members; within each group, I collected feeding data for the dominant adult male, the highest ranking pregnant female, and the highest ranking female with a young infant. There were no significant differences between groups during the study in either the mean estimated energy value of the food ingested per day for each individual (385±27 kJ kg^-1 day^-1) or in the estimated energy expended to obtain that food (114±3 kJ kg^-1 day^-1). Mean foraging efficiency ratios, which reflect net energy gain per unit of foraging time, also did not vary as a function of the size of the group in which the baboons were living. There was substantial variation between days in the efficiency ratios of all animals; this was the result of large differences in energy intake rather than in the energy expended during foraging itself. The members of the smallest group spent on the average only one-half as much time feeding each day as did individuals in the two larger groups. However, they obtained almost as much energy while foraging, primarily because their rate of food intake while actually eating tended to be higher than the rate in the other groups. The baboons in the small group were observed closer to trees that they could climb to escape ground predators, and they also were more likely to sit in locations elevated above the ground while resting. Such differences would be expected if the members of the small group were less able to detect approaching predators than individuals that lived in the larger groups. The results of this study suggest that predator detection or avoidance, rather than increased foraging efficiency, may be the primary benefit of living in larger groups in this population.     

Chick-a-dee call variation in the context of “flying” avian predator stimuli: a field study of Carolina chickadees (<Emphasis Type="Italic">Poecile carolinensis</Emphasis>)

Chick-a-dee calls function in social organization in Poecile (chickadee) species. Recent field and aviary studies have found that variation in chick-a-dee calls relates to the type or proximity of avian predator, or level of threat. Earlier studies on calls in the context of predator stimuli have typically used stationary and perched predator models. For chickadees and other small songbirds, more frequently detected and more dangerous avian predatory stimuli are flying predators. In the present study, we tested whether simulated flying avian predator and control models influenced chick-a-dee calling behavior of wild Carolina chickadees, Poecile carolinensis. At 20 independent field sites, chickadee subjects were presented with wooden models that were painted to resemble either a predatory sharp-shinned hawk (Accipiter striatus) or a blue jay (Cyanocitta cristata) and that were made to “fly” down a zip line near a feeding station chickadees were using. The note composition of chick-a-dee calls was affected by both the flight of stimuli and type of model. Call variation in this flying predator context suggests interesting similarities and differences with experimental findings with congeners. Finally, increased production of certain notes to the flying of both model types provides support for a “Better Safe than Sorry” strategy. When costs of alarm calling are low but costs of discriminating potentially serious threats may be extremely high, individuals should err on the side of caution, and alarm call to any potentially threatening stimulus.     

Exploring the effects of individual traits and within-colony variation on task differentiation and collective behavior in a desert social spider

Social animals are extraordinarily diverse and ecologically abundant. In understanding the success of complex animal societies, task differentiation has been identified as a central mechanism underlying the emergence and performance of adaptive collective behaviors. In this study, we explore how individual differences in behavior and body size determine task allocation in the social spider Stegodyphus dumicola. We found that individuals with high body condition indices were less likely to participate in prey capture, and individuals’ tendency to engage in prey capture was not associated with either their behavioral traits or body size. No traits were associated with individuals’ propensity to participation in web repair, but small individuals were more likely to engage in standard web-building. We also discovered consistent, differences among colonies in their collective behavior (i.e., colony-level personality). At the colony level, within-colony variation in behavior (aggressiveness) and body size were positively associated with aggressive foraging behavior. Together, our findings reveal a subtly complex relationship between individual variation and collective behavior in this species. We close by comparing the relationship between individual variation and social organization in nine species of social spider. We conclude that intraspecific variation is a major force behind the social organization of multiple independently derived lineages of social spider.     

The social network structure of a wild meerkat population: 2. Intragroup interactions

Knowledge of the structure of networks of social interactions is important for understanding the evolution of cooperation, transmission of disease, and patterns of social learning, yet little is known of how environmental, ecological, or behavioural factors relate to such structures within groups. We observed grooming, dominance, and foraging competition interactions in eight groups of wild meerkats (Suricata suricatta) and constructed interaction networks for each behaviour. We investigated relationships between networks for different social interactions and explored how group attributes (size and sex ratio), individual attributes (tenure of dominants), and ecological factors (ectoparasite load) are related to variation in network structure. Network structures varied within a group according to interaction type. Further, network structure varied predictably with group attributes, individual attributes, and ecological factors. Networks became less dense as group size increased suggesting that individuals were limited in their number of partners. Groups with more established dominant females were more egalitarian in their grooming and foraging competition interactions, but more despotic in their dominance interactions. The distribution of individuals receiving grooming became more skewed at higher parasite loads, but more equitable at low parasite loads. We conclude that the pattern of interactions between members of meerkat groups is not consistent between groups but instead depends on general attributes of the group, the influence of specific individuals within the group, and ecological factors acting on group members. We suggest that the variation observed in interaction patterns between members of meerkat groups may have fitness consequences both for individual group members and the group itself.     

Foraging behavior of a generalist marine top predator,Japanese cormorants (<Emphasis Type="Italic">Phalacrocorax filamentosus</Emphasis>), in years of demersal versus epipelagic prey

To examine the behavioral adjustment of a generalist marine top predator to variability of their prey, we studied the foraging behavior of Japanese cormorants (Phalacrocorax filamentosus) breeding at Teuri Island, Hokkaido, in years of contrasting demersal and epipelagic prey composition. We used radio telemetry and ship-based surveys to determine behavior and at-sea distribution during three summers (1996–1998). The cormorants fed on epipelagic anchovy (Engraulis japonicus) and sandlance (Ammodytes personatus) in 1998 (year of epipelagic diet), while they fed on benthic rock fish (Sebastes spp.) and flatfish (Pleuronectidae) and nearshore-living naked sandlance (Hypophychus dybowskii), as well as epibenthic greenling (Hexagrammidae) in 1996 and 1997 (year of demersal diet). Cormorants engaged in larger feeding groups, visited more feeding sites, and stayed at each feeding site for a shorter period in the year of epipelagic diet than in the years of demersal diet. The cormorants made long foraging trips and fed in the mainland coastal habitat, distant from the colony, in the years of demersal diet. Individual radio-tracked birds fed over the wide area between the islands and mainland, in the year of epipelagic diet, while most individuals specialized in mainland or island coastal habitats in the years of demersal diet. Behavioral adjustment of Japanese cormorants might allow them to exploit both unpredictable epipelagic and predictable benthic prey efficiently.Communicated by T. Ikeda, Hakodate     

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.     

Colony state and regulation of pollen foraging in the honey bee,Apis mellifera L.

Summary To place social insect foraging behavior within an evolutionary context, it is necessary to establish relationships between individual foraging decisions and parameters influencing colony fitness. To address this problem, we examined interactions between individual foraging behavior and pollen storage levels in the honey bee, Apis mellifera L. Colonies responded to low pollen storage conditions by increasing pollen intake rates 54% relative to high pollen storage conditions, demonstrating a direct relationship between pollen storage levels and foraging effort. Approximately 80% of the difference in pollen intake rates was accounted for by variation in individual foraging effort, via changes in foraging activity and individual pollen load size. An additional 20% resulted from changes in the proportion of the foraging population collecting pollen. Under both high and low pollen storage treatments, colonies returned pollen storage levels to pre-experimental levels within 16 days, suggesting that honey bees regulate pollen storage levels around a homeostatic set point. We also found a direct relationship between pollen storage levels and colony brood production, demonstrating the potential for cumulative changes in individual foraging decisions to affect colony fitness. Offprint requests to: J.H. Fewell at the current address     

Divergent foraging strategies of three co-occurring north Pacific flatfishes

Despite facing similar constraints imposed by the environment, significant variation in life history traits frequently exists among species generally considered to comprise a single ecological guild. For juvenile flatfishes, constraints on foraging activity include variation in light and prey availability, as well as predation risk. This paper describes the visual constraints on, and divergent foraging strategies of three co-occurring north Pacific flatfish species, northern rock sole (Lepidopsetta polyxystra), Pacific halibut (Hippoglossus stenolepis), and English sole (Pleuronectes vetulus). Visual foraging abilities measured in the laboratory decreased rapidly below 10⁻⁴ μmol photons·m⁻² s⁻¹, and were similar among species. Despite similar sensory constraints, field sampling in August 2004 at a Kodiak Island nursery site (Holiday Beach, 57^o41.2′ N, 152^o27.7′ W) identified species differences in diets, diel foraging patterns, and within-nursery depth distributions. Northern rock sole and English sole fed primarily on bivalve siphons and polychaetes, whereas mysids dominated the diets of Pacific halibut. Northern rock sole were geographically the most widespread but feeding activity was temporally restricted to the dusk period. Pacific halibut were rare in shallow depths (<5 m) and fed most intensively prior to dusk. English sole fed throughout the daylight hours and were abundant only in the shallowest (<5 m) habitats. These differences in diets, foraging times, and habitat use appear related to previously documented species-specific behavioral characteristics as well as general spatial (increasing with depth) and temporal (increasing during foraging activity) variations in predation risk. At one extreme, the conservative behavioral strategy of northern rock sole may permit use of a broader range of foraging habitats, whereas English sole may be restricted to shallow water by limited behavioral responses to predation threat. These observations demonstrate that the appearance of habitat partitioning is not due to differences in sensory ability, but reflects multi-faceted, species-specific responses to the ecological tradeoffs between foraging and predation risks.     

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     

Family-related differences in social foraging tactic use in the zebra finch (Taeniopygia guttata)

When animals forage in groups, they can search for food themselves (producer tactic), or they can search for opportunities to exploit the food discoveries of others (scrounger tactic). Both theoretical and empirical work have shown that group-level use of these alternative tactics is influenced by environmental conditions including group size and food distribution, and individual tactic use can be influenced by several measures of individual state, including body condition. Because body condition has been shown to be heritable for various species, social foraging tactics may also be heritable. We looked for evidence of heritability in social foraging tactic use in the zebra finch (Taeniopygia guttata) by testing whether: (1) natural variation in body condition correlates with tactic use, (2) there are family-related differences in body condition, and (3) there are family-related differences in observed tactic use. Tactic use in the zebra finch was significantly related to body condition; individuals with lower body condition scores had a significantly higher use of the scrounger tactic as predicted from variance-sensitive producer–scrounger models. Body-condition scores differed significantly between families, suggesting that this aspect of individual state may have a heritable component. Finally, we recorded significant family-related differences in the use of producer and scrounger alternatives. These results are consistent with heritability in observed tactic use resulting from an inheritance of individual state, in this case body condition, which itself influences tactic use. Understanding how and why individuals differ in their use of alternative tactics is fundamental as it may provide important insights into inter-individual variation in fitness.