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Although ecological disturbances can have a strong influence on pollinators through changes in habitat, virtually no studies have quantified how characteristics of wildfire influence the demography of essential pollinators. Nevertheless, evaluating this topic is critical for understanding how wildfire is linked to pollinator population dynamics, particularly given recent changes in wildfire frequency and severity in many regions of the world. In this study, we measured the demographic response of the blue orchard bee (Osmia lignaria) across a natural gradient of wildfire severity to assess how variation in wildfire characteristics influenced reproductive output, offspring sex ratio, and offspring mass. We placed nest blocks with a standardized number and sex ratio of pre-emergent adult bees across the wildfire gradient, finding some evidence for a positive but highly variable relationship between reproductive output and fire severity surrounding the nest site at both local (100 m) and landscape (750 m) scales. In addition, the production of female offspring was > 10% greater at nest sites experiencing the greatest landscape-scale fire severity relative to the lowest-severity areas. The finding that blue orchard bees biased offspring production towards the more expensive offspring sex with increasing fire severity shows a functional response to changes in habitat quality through increased density of flowering plants. Our findings indicate that burned mixed-conifer forest provides forage for the blue orchard bee across a severity gradient, and that the increase in floral resources that follows high-severity fire leads females to shift resource allocation to the more costly sex when nesting.

The extent to which migratory bats forage at stopover sites or while in migratory flight is poorly understood. Endogenous fat stores have lower δ13C values relative to the dietary substrates from which they were synthesized, and so, the fed versus fasted state of bats should be discernable by comparing their breath δ13C at capture to that after a known period of fasting. We captured silver-haired bats (Lasionycteris noctivagans) at a stopover site at Long Point, Ontario, Canada, during spring and fall migration. We collected breath samples at capture and after fasting in captivity for 12 h, providing a fasted-state δ13C value corresponding to metabolism of fat stores. We also collected and weighed fecal pellets produced while in captivity. Breath δ13C values at capture were positively correlated with mass of feces produced. During spring migration, δ13C values of breath CO2 at capture were low and similar to fasting values, but increased with date consistent with increased foraging at stopover and reliance on exogenous dietary nutrients as the season progressed. The opposite temporal pattern was found during fall migration. Our findings suggest that bats forage during migratory stopover when environmental conditions permit despite potential time trade-offs between feeding and travel, and the energy savings resulting from torpor during roosting. This study provides insight into the eco-physiology of bat migration and shows the importance of foraging habitat for migratory bats.

Life history theory predicts trade-offs in allocation between survival, maintenance, growth, and reproduction, especially when resources are scarce. Individual variation in resource acquisition can affect trade-offs, but is often unaccounted for. We quantified the fitness costs of reproduction, accounting for environmental conditions, maternal characteristics and individual variation. We analyzed 10 years of data from marked kangaroos to evaluate how reproductive allocation affected annual mass change and skeletal growth, subsequent fecundity and weaning success, and survival, accounting for maternal mass or size and forage availability. Through repeated measurements of 76–91 females, we investigated how trade-offs varied within and between individuals, assessing whether individual variation could mask population-level trade-offs. In poor environments, females that weaned an offspring lost mass. Females that nursed an offspring for > 7 months had reduced skeletal growth. Females that did not gain mass over the previous 12 months rarely reproduced, especially if they had nursed an offspring for > 7 months the previous year. Reproductive allocation had no effect on weaning success, which was very low, and did not affect maternal survival, suggesting a conservative strategy. Disentangling within- and between-individual responses revealed trade-offs within individuals, but because individuals did not vary in their responses to earlier effort, these trade-offs did not drive population trends. The interacting effects of environmental conditions, maternal characteristics and individual variation on allocation trade-offs demonstrate the importance of long-term monitoring for understanding life history variations in changing environments.

The landscape of fear is an important driver of prey space use. However, prey can navigate the landscape of fear by exploiting temporal refuges from predation risk. We hypothesized that diel patterns of predator and prey movement and space use would be inversely correlated due to temporal constraints on predator habitat domain. Specifically, we evaluated habitat selection and activity of the vicuña and its only predator, the puma, during three diel periods: day, dawn/dusk, and night. Pumas selected the same habitats regardless of diel period-vegetated and rugged areas that feature stalking cover for pumas-but increased their activity levels during dawn/dusk and night when they benefit from reduced detection by prey. Vicuñas avoided areas selected by pumas and reduced activity at night, but selected vegetated areas and increased activity by day and dawn/dusk. Vicuña habitat selection and movement strategies appeared to reduce the risk of encountering pumas; movement rates of pumas and vicuñas were negatively correlated across the diel cycle, and habitat selection was negatively correlated during dawn/dusk and night. Our study shows that an ambush predator's temporal activity and space use patterns interact to create diel refugia and shape the antipredator behaviors of its prey. Importantly, it is likely the very nature of ambush predators' static habitat specificity that makes predator activity important to temporally varying perceptions of risk. Prey which depend on risky habitats for foraging appear to mitigate risk by feeding when they can more easily detect predators and when predators are least active.

Sufficiently diverse and abundant resources are essential for generalist consumers, and form an important part of a suite of conservation strategies for pollinators. Honey bees are generalist foragers and are dependent on diverse forage to adequately meet their nutritional needs. Through analysis of stored pollen (bee bread) samples obtained from 26 honey bee (Apis mellifera L.) hives across NW-England, we quantified bee bread nutritional content and the plant species that produced these stores from pollen. Protein was the most abundant nutrient by mass (63%), followed by carbohydrates (26%). Protein and lipid content (but not carbohydrate) contributed significantly to ordinations of floral diversity, linking dietary quality with forage composition. DNA sequencing of the ITS2 region of the nuclear ribosomal DNA gene identified pollen from 89 distinct plant genera, with each bee bread sample containing between 6 and 35 pollen types. Dominant genera included dandelion (Taraxacum), which was positively correlated with bee bread protein content, and cherry (Prunus), which was negatively correlated with the amount of protein. In addition, proportions of amino acids (e.g. histidine and valine) varied as a function of floral species composition. These results also quantify the effects of individual plant genera on the nutrition of honey bees. We conclude that pollens of different plants act synergistically to influence host nutrition; the pollen diversity of bee bread is linked to its nutrient content. Diverse environments compensate for the loss of individual forage plants, and diversity loss may, therefore, destabilize consumer communities due to restricted access to alternative resources. Electronic supplementary material The online version of this article (10.1007/s00442-017-3968-3) contains supplementary material, which is available to authorized users.

The seasonal movement of animals at broad spatial scales provides insight into life-history, ecology and conservation. By combining high-resolution satellite-tagged data with hierarchical Bayesian movement models, we can associate spatial patterns of movement with marine animal behavior. We used a multi-state mixture model to describe humpback whale traveling and area-restricted search states as they forage along the West Antarctic Peninsula. We estimated the change in the geography, composition and characteristics of these behavioral states through time. We show that whales later in the austral fall spent more time in movements associated with foraging, traveled at lower speeds between foraging areas, and shifted their distribution northward and inshore. Seasonal changes in movement are likely due to a combination of sea ice advance and regional shifts in the primary prey source. Our study is a step towards dynamic movement models in the marine environment at broad scales.

Animals may partially overcome environmental constraints on fitness by behaviorally adjusting their exposure to costs and supplies of energy. Few studies, however, have linked spatiotemporal variation in the energy landscape to behaviorally mediated measures of performance that ostensibly influence individual fitness. We hypothesized that strength of selection by North American elk (Cervus elaphus) for areas that reduced costs of thermoregulation and activity, and increased access to high-quality forage, would influence four energetically mediated traits related to fitness: birth mass of young, nutritional condition of adult females at the onset of winter, change in nutritional condition of females between spring and winter, and neonatal survival. We used a biophysical model to map spatiotemporally explicit costs of thermoregulation and activity experienced by elk in a heterogeneous landscape. We then combined model predictions with data on forage characteristics, animal locations, nutritional condition, and mass and survival of young to evaluate behaviorally mediated effects of the energy landscape on fitness-related traits. During spring, when high-quality forage was abundant, female elk that consistently selected low-cost areas before parturition gave birth to larger young than less-selective individuals, and birth mass had a strong, positive influence on probability of survival. As forage quality declined during autumn, however, lactating females that consistently selected the highest quality forage available accrued more fat and entered winter in better condition than less-selective individuals. Results of our study highlight the importance of understanding the dynamic nature of energy landscapes experienced by free-ranging animals.

Fission–fusion dynamics hypothetically enable animals to exploit dispersed and ephemeral food resources while minimizing predation risk. Disentangling factors affecting group size and composition of fission–fusion species facilitates their management and conservation. We used a 6-year data set of 2888 group formations of Masai giraffes in Tanzania to investigate determinants of social group size and structure. We tested whether ecological (lion density, vegetation structure, and prevalence of primary forage plants), anthropogenic (proximity to human settlements), temporal (rainy or dry season), and social (local giraffe density, adult sex ratio, and proportion of calves) factors explained variation in group size and sex- and age-class composition. Food availability rather than predation risk mediated grouping dynamics of adult giraffes, while predation risk was the most important factor influencing congregations with calves. Smallest group sizes occurred during the food-limiting dry season. Where predation risk was greatest, groups with calves were in bushlands more than in open grasslands, but the groups were smaller in size, suggesting mothers adopted a strategy of hiding calves rather than a predator-detection-and-dilution strategy. Groups with calves also were farther from towns but closer to traditional human compounds (bomas). This may be due to lower predator densities, and thus reduced calf predation risk, near bomas but higher human disturbance near towns. Sex- and age-based differences in habitat use reflected nursing mothers’ need for high-quality forage while also protecting their young from predation. Our results have implications for conservation and management of giraffes and other large-bodied, herd-forming ungulates in heterogeneous environments subject to anthropogenic threats.

The temporal distribution of resources is an important aspect of habitat quality that can substantially impact population success. Although it is widely accepted that floral resources directly influence wild bee population sizes, we lack experimental data evaluating how resource availability affects colony growth via demographic mechanisms. To achieve this, we tracked marked individuals in bumble bee (Bombus vosnesenskii) colonies to evaluate whether worker survival and reproduction responded to experimentally elevated forage early in colony development. Specifically, we assessed the effect of early resource environment on worker and sexual offspring production, and the survival and body size of individual workers. We also assessed whether responses of colonies differed when exposed to higher or lower resource environments at a relatively smaller (~ 10 workers) or larger (~ 20 workers) size. Resource supplementation always resulted in greater total offspring and male production; however, the influence of supplementation on worker production and quality depended on colony size at the start of supplementation. Among colonies that were initially smaller, colonies that were supplemented produced fewer but larger bodied and longer lived workers compared to control counterparts. Among colonies that were initially larger, colonies that were supplemented produced more workers than corresponding controls, but without changes to worker quality. Collectively, these results provide clear experimental evidence that greater resource availability early in colony development increases overall productivity, and indicate that colonies may pursue different allocation strategies in response to the resource environment, investing in more or better workers.

Bumble bee abundance in agricultural landscapes is known to decrease with increasing distance from seminatural grasslands, but whether the pollination of bumble-bee-pollinated wild plants shows a similar pattern is less well known. In addition, the relative effects of landscape composition (landscape heterogeneity) and landscape configuration (distance from seminatural grassland) on wild plant pollination, and the interaction between these landscape effects, have not been studied using landscape-level replication. We performed a field experiment to disentangle these landscape effects on the pollination of a native herb, the sticky catchfly (Lychnis viscaria), while accounting for the proportion of oilseed rape across landscapes and the local abundance of bee forage flowers. We measured pollen limitation (the degree to which seed set is pollen-limited), seed set, and seed set stability using potted plants placed in landscapes that differed in heterogeneity (composition) and distance from seminatural grassland (configuration). Pollen limitation and seed set in individual plants did not respond to landscape composition, landscape configuration, or proportion of oilseed rape. Instead, seed set increased with increasing local bee forage flower cover. However, we found within-plant variability in pollen limitation and seed set to increase with increasing distance from seminatural pasture. Our results suggest that average within-plant levels of pollen limitation and seed set respond less swiftly than the within-plant variability in pollen limitation and seed set to changes in landscape configuration. Although landscape effects on pollination were less important than predicted, we conclude that landscape configuration and local habitat characteristics play larger roles than landscape composition in the pollination of L. viscaria.

The giving-up density framework is an elegant and widely adopted mathematical approach to measuring animals' foraging decisions at non-replenishing artificial resource patches. Under this framework, an animal should "give up" when the benefits of foraging are outweighed by the costs (e.g., predation risk, energetic, and/or missed opportunity costs). However, animals of many species may forage in groups, and group size is expected to alter perceived predation risk and hence influence quitting decisions. Yet, most giving-up density studies assume either that individuals forage alone or that giving-up densities are not affected by group foraging. For animals that forage both alone and in groups, differences in giving-up densities due to group foraging rather than experimental variables may substantially alter interpretation. However, no research to date has directly investigated how group foraging affects the giving-up density. We used remote-sensing cameras to identify instances of group foraging in two species of Rattus across three giving-up density experiments to determine whether group foraging influences giving-up densities. Both Rattus species have been observed to vary between foraging alone and in groups. In all three experiments, solo foragers left higher giving-up densities on average than did group foragers. This result has important implications for studies using giving-up densities to investigate perceived risk, the energetic costs of searching, handling time, digestion, and missed opportunity costs, particularly if groups of animals are more likely to experience certain experimental treatments. It is critically important that future giving-up density studies consider the effects of group foraging.

Pollinator behavior influences plant reproduction in many ways. A traditional measure of pollination, the number of visits received, may be a poor predictor of plant reproductive success, particularly when there are trade-offs between visit quantity and components of visit quality. For example, the duration of pollinator visits may be negatively correlated with the number of visits received by a flower. We tested for a trade-off between the number of bumblebee visits and the duration of those visits in an experimental population of snapdragons (Antirrhinummajus: Scrophulariaceae). The duration of a bumblebee visit to a flower increased significantly with the time interval since the flower had last been visited. Over the lifetime of a flower the correlation between the total number and average duration of visits received by a flower was weakly negative. However, at the whole-plant level the correlation was positive: plants whose flowers received more visits also received visits of longer duration. Factors affecting the relationship between quantity and duration of pollinator visits to flowers also were investigated. Two factors weakened the negative dependence of average visit duration on number of visits received by individual snapdragon flowers: (1) the correlation between the total number of visits to a flower and the average interval between visits was only −0.53, as visits to individual flowers were not very evenly spaced over time, and (2) newly opened flowers received fewer and shorter visits than older flowers. Comparing whole plants, nectar production per flower varied dramatically across individuals, a probable explanation for the positive correlation between visit number and average duration per flower observed at the plant level. The potential for a trade-off between these two components of pollinator service exists when visit duration depends on reward quantity; whether the trade-off is realized will depend on variation in nectar production and on whether pollinators forage systematically.

Optimality models of food selection by herbivores assume that individuals are capable of assessing forage value, either directly through the currency used in the model or indirectly through other variables correlated with the currency. Although energy and protein are the two currencies most often used, controversy exists regarding their respective influence on food choice. Part of the debate is due to the difficulty of teasing apart these two nutrients, which are closely correlated in most natural forages. Here we offer a test of the assumption that energy and protein contents of the forage are both currencies that large mammalian herbivores can use when selecting their food. We observed feeding behavior of 47 wild white-tailed deer (Odocoileusvirginianus) during winter while individuals were presented with four experimental foods representing two levels of energy and protein (dry matter digestibility: 40–50%; crude protein: 12–16%). Using experimental foods allowed us to separate the influences of energy and protein and clearly distinguish between the roles of these two nutrients. Deer discriminated between foods through partial selection, and selected diets higher in energy but lower in protein. The observed choices appeared consistent with physiological needs of deer wintering at the study site, where digestible energy was in short supply in the natural environment while protein was probably not. Results are in good agreement with recent findings on domesticated ruminants. They support a basic assumption of optimality models of food selection that use energy and/or protein as a currency, although the physiological mechanisms behind the food selection process remain unclear. We urge students of food selection by herbivores to replicate our experiment with other foods and/or in other circumstances before more general conclusions are drawn.

We test predictions about differences in the foraging behaviors of male and female giraffes (Giraffa camelopardalis tippelskirchi Matchie) that derive from a hypothesis linking sexual size dimorphism to foraging behavior. This body-size hypothesis predicts that males will exhibit specific behaviors that increase their dry-matter intake rate relative to females. Foraging behavior was examined at two hierarchical levels corresponding to two spatial and temporal scales, within patches and within habitats. Patches are defined as individual trees or shrubs and habitats are defined as collections of patches within plant communities. Males were predicted to increase dry-matter intake rate within patches by taking larger bites, cropping bites more quickly, chewing less, and chewing faster. Within habitats, males were expected to increase intake rate by increasing the proportion of foraging time devoted to food ingestion as opposed to inter-patch travel time and vigilance. The predictions were tested in a free-ranging population of giraffes in Mikumi National Park, Tanzania. Males spent less total time foraging than females but allocated a greater proportion of their foraging time to forage ingestion as opposed to travel between patches. There was no sex difference in rumination time but males spent more time in activities other than foraging and rumination, such as walking. Within patches, males took larger bites than females, but females cropped bites more quickly and chewed faster. Males had longer per-bite handling times than females but had shorter handling times per gram of intake. Within habitats, males had longer average patch residence times but there was no significant sex difference in inter-patch travel times. There was no overall difference between sexes in vigilance while foraging, although there were significant sex by habitat and sex by season interactions. Although not all the predictions were confirmed, overall the results agree qualitatively with the body-size hypothesis. Sex-related differences in foraging behavior led to greater estimated intake rates for males at the within-patch and within-habitat scales.

Recent theoretical models have considered how nutritional status and predation risk affect the decision by an individual to forage. We consider the feeding behaviour of a scavenger under risk of predation. The intertidal gastropod Nassarius dorsatus Roding was used to test the following hypotheses: (1) a damaged conspecific should be a less preferred meal than a damaged predator or another species; (2) a scavenger should be able to discriminate between a damaged conspecific in the presence and one in the absence of its predator, preferring the latter; and (3) the decision by a scavenger to feed should reflect a trade-off between nutritional status and the above preferences. Results from the laboratory and field were consistent with predictions. Two hypotheses are suggested from the work: scavengers may generally be less willing to feed on carcases of conspecifics than on those of other species, and intertidal scavengers may commonly face a relative or absolute shortage of food.

Interactions among granivores and seeds depend on the foraging behaviour and morphology of the granivores and on the attributes and availability of seeds. We investigated seed selection by the seed harvesting ant Messor bouvieri in three adjacent plant communities in Spain by relating the harvested seeds to those in the seed rain. Preference was positively correlated with seed size and abundance which accounted for 43% and 20% of the variance respectively. Contrary to predictions of central place foraging theory, the size of seeds harvested did not increase with distance from the nest. Inclusion of a less-preferred item in the diet was more strongly related to the abundance of more-preferred items (60% of the variance) than the abundance of the less-preferred item (14% of the variance). Worker size accounted for 20–30% of the variance in the size of harvested seeds, although small workers did not appear to be constrained by load size for the range of seeds available. The body size of ants was significantly larger in the community with the greatest proportion of large seeds, although this was not due to their ability to carry larger loads or due to the greater force required to crush these seeds. The strong preference of M. bouvieri for large seeds may have important consequences for the plant communities in which they forage.

In non-territorial species, individuals can move freely and should be distributed in an ideal free manner between habitats and areas with respect to resources that influence lifetime reproductive success (LRS). Consequently, no relationship between diet quality and LRS should be expected. However, there have been no attempts to test this prediction. The present paper investigates the relationship between forage habitat use and LRS in red deer (Cervus elaphus) hinds within three neighbouring areas on the Isle of Rum which differed in their amounts of high-quality-forage habitat. Within areas, hinds move widely and have access to the same resources. We found no correlation between LRS of individual hinds and their use of high-quality-forage habitat (i.e. short Agrostis/Festuca grassland). Our analysis suggests that high hind densities on short Agrostis/Festuca grassland offset any advantages of increased access to preferred forage. These results support the hypothesis that red deer hinds are distributed in an ideal free manner with respect to the use of high-quality-forage habitat. However, hinds rarely leave areas where they are born, and the analysis suggests that constraints in changing areas hindered an ideal free distribution on a larger spatial scale. Consequently, mean LRS was not the same within the three investigated areas: one area, with a low amount of short Agrostis/Festuca grassland and a low hind density, contributed more male offspring (and more total offspring) per hind to the population than the other two areas.

Both atmospheric [CO2] and average surface temperatures are predicted to increase with potentially different, additive or opposing, effects on leaf quality and insect herbivore activity. Few studies have directly measured the interactive effects of concurrent changes in [CO2] and temperature on insect herbivores. None have done so over the entire developmental period of a tree-feeding insect, and none have compared responses to low pre-industrial [CO2] and present day [CO2] to estimate responses to future increases. Eucalypt herbivores may be particularly sensitive to climate-driven shifts in plant chemistry, as eucalypt foliage is naturally low in [N]. In this study, we assessed the development of the eucalypt herbivore Doratifera quadriguttata exposed concurrently to variable [CO2] (290, 400, 650 μmol mol(-1)) and temperature (ambient, ambient +4 °C) on glasshouse-grown Eucalyptus tereticornis. Overall, insects performed best on foliage grown at pre-industrial [CO2], indicating that modern insect herbivores have already experienced nutritional shifts since industrialisation. Rising [CO2] increased specific leaf mass and leaf carbohydrate concentration, subsequently reducing leaf [N]. Lower leaf [N] induced compensatory feeding and impeded insect performance, particularly by prolonging larval development. Importantly, elevated temperature dampened the negative effects of rising [CO2] on larval performance. Therefore, rising [CO2] over the past 200 years may have reduced forage quality for eucalypt insects, but concurrent temperature increases may have partially compensated for this, and may continue to do so in the future. These results highlight the importance of assessing plant-insect interactions within the context of multiple climate-change factors because of the interactive and potentially opposing effects of different factors within and between trophic levels.

Herbivores are usually distributed unevenly across the landscape often because of variation in resource availability. We used zero-inflated generalised additive models (to account for data with a high number of zeros) that include georeferences to predict winter distribution of a large herbivore (moose Alces alces). Moose distribution was analysed in relation to forage availability and distance to neighbouring sites. Our results showed that the ability to explain moose distribution indexed by pellet count data at a local scale increased when spatial information (longitude and latitude) was added to the model compared to the model only including food availability. By using the relationship between moose and forage distribution, and the spatial information, we predicted patch choice by moose reasonably well in 2 out of 4 years. However, the distribution of moose was also influenced by weather conditions, as it was most clumped in the year with most snow. In conclusion, our study lends support for a non-linear approach using georeferences for a comprehensive understanding of herbivore distribution at a small scale. This result also indicates that the use of a certain patch by moose not only depends on the selected patch itself but is also influenced by the neighbouring patch and factors at a larger spatial scale, such as moose management influencing the density above moose home range level. The relatively high proportion of unexplained variation suggests that the use of a certain patch is also influenced by other factors such as topography, predation, competition, weather conditions, and wildlife management strategies.

In habitats where prey is either rare or difficult to predict spatiotemporally, such as open habitats, predators must be adapted to react effectively to variations in prey abundance. Open-habitat foraging bats have a wing morphology adapted for covering long distances, possibly use information transfer to locate patches of high prey abundance, and would therefore be expected to show an aggregative response at these patches. Here, we examined the effects of prey abundance on foraging activities of open-habitat foragers in comparison to that of edge-habitat foragers and closed-habitat foragers. Bat activity was estimated by counting foraging calls recorded with bat call recorders (38,371 calls). Prey abundance was estimated concurrently at each site using light and pitfall traps. The habitat was characterized by terrestrial laser scanning. Prey abundance increased with vegetation density. As expected, recordings of open-habitat foragers clearly decreased with increasing vegetation density. The foraging activity of edge- and closed-habitat foragers was not significantly affected by the vegetation density, i.e., these guilds were able to forage from open habitats to habitats with dense vegetation. Only open-habitat foragers displayed a significant and proportional aggregative response to increasing prey abundance. Our results suggest that adaptations for effective and low-cost foraging constrains habitat use and excludes the guild of open-habitat foragers from foraging in habitats with high prey abundance, such as dense forest stands.

Anuran amphibians have traditionally been considered suitable model organisms for community ecology studies. In this paper, we review and quantitatively re-analyze the original studies published on dietary relationships among anuran communities worldwide. We re-analyzed 33 independent communities from all continents, including data on prey numbers and/or prey volumes. All datasets underwent null model re-analysis using two randomization algorithms. In addition, logistic regression models were used to identify specific assemblage patterns. We discovered three main patterns: (1) one particular algorithm (RA2, which replaces every cell in the matrix with one randomly chosen, but retains the zero structure of the matrix) performed much better at uncovering community structure; (2) nonrandom structure was more likely to occur in tropical than in nontropical regions; (3) prey volume was a better descriptor of dietary relationships than prey number when attempting to detect nonrandom structure. We showed that: pattern (1) depended on both the overall generalist nature of the anurans in general and on the mathematical properties of RA2; pattern (2) was consistent with theoretical attributes of tropicality; and pattern (3) was due to the intrinsic ecological properties of generalist organisms, which forage more or less opportunistically on prey of very different sizes; hence, when prey number is used as a diet descriptor, there is a risk of approximating different prey taxa that make different relative volume contributions to the overall diet as being of similar relevance .

Identifying factors shaping variation in resource selection is central for our understanding of the behaviour and distribution of animals. We examined summer habitat selection and space use by 108 Global Positioning System (GPS)-collared moose in Norway in relation to sex, reproductive status, habitat quality, and availability. Moose selected habitat types based on a combination of forage quality and availability of suitable habitat types. Selection of protective cover was strongest for reproducing females, likely reflecting the need to protect young. Males showed strong selection for habitat types with high quality forage, possibly due to higher energy requirements. Selection for preferred habitat types providing food and cover was a positive function of their availability within home ranges (i.e. not proportional use) indicating functional response in habitat selection. This relationship was not found for unproductive habitat types. Moreover, home ranges with high cover of unproductive habitat types were larger, and smaller home ranges contained higher proportions of the most preferred habitat type. The distribution of moose within the study area was partly related to the distribution of different habitat types. Our study shows how distribution and availability of habitat types providing cover and high-quality food shape ungulate habitat selection and space use. Electronic supplementary material The online version of this article (doi:10.1007/s00442-011-2072-3) contains supplementary material, which is available to authorised users.

Arctic Cervids face considerable challenges in sustaining life in a harsh and highly seasonal environment, and when to forage is a key component of the survival strategy. We predict that a cervid maximizes net intake of energy to change the duration of feeding-ruminating cycles depending on season, and pays no attention to light or other activity-entraining cues. Still, in periods of bad weather it may pay energetically to reduce exposure and heat loss. We investigated environmental impact on the seasonal and daily activity pattern of a food-limited, predator-free arctic deer, the Svalbard reindeer. We found that the reindeer indeed had season-dependent feeding-rumination intervals, with no distinct peaks in activity at sunrise and sunset, as would be expected if animals maximize energy intake rates in predator-free environments. However, they temporarily reduced activity when exposed to low temperature and increased precipitation during winter, possibly to conserve energy. We provide insight into the behavioural strategy of Svalbard reindeer which enables them to cope with such an extreme environment.

Avoiding predators most often entails a food cost. For the Scandinavian brown bear (Ursus arctos), the hunting season coincides with the period of hyperphagia. Hunting mortality risk is not uniformly distributed throughout the day, but peaks in the early morning hours. As bears must increase mass for winter survival, they should be sensitive to temporal allocation of antipredator responses to periods of highest risk. We expected bears to reduce foraging activity at the expense of food intake in the morning hours when risk was high, but not in the afternoon, when risk was low. We used fine-scale GPS-derived activity patterns during the 2 weeks before and after the onset of the annual bear hunting season. At locations of probable foraging, we assessed abundance and sugar content, of bilberry (Vaccinium myrtillus), the most important autumn food resource for bears in this area. Bears decreased their foraging activity in the morning hours of the hunting season. Likewise, they foraged less efficiently and on poorer quality berries in the morning. Neither of our foraging measures were affected by hunting in the afternoon foraging bout, indicating that bears did not allocate antipredator behavior to times of comparably lower risk. Bears effectively responded to variation in risk on the scale of hours. This entailed a measurable foraging cost. The additive effect of reduced foraging activity, reduced forage intake, and lower quality food may result in poorer body condition upon den entry and may ultimately reduce reproductive success. Electronic supplementary material The online version of this article (doi:10.1007/s00442-016-3729-8) contains supplementary material, which is available to authorized users.

Many bumblebee species are declining at a rapid rate in the United Kingdom and elsewhere. This is commonly attributed to the decline in floral resources that has resulted from an intensification in farming practices. Here we assess growth of nests of the bumblebee, Bombus terrestris, in habitats providing different levels of floral resources. Experimental nests were placed out in conventional farmland, in farmland with flower-rich conservation measures and in suburban areas. Nests gained weight more quickly and attained a larger final size in suburban areas compared to elsewhere. The diversity of pollens gathered by bees was highest in suburban areas, and lowest in conventional farmland. Nests in suburban areas were also more prone to attack by the specialist bumblebee parasite Aphomia sociella, suggesting that this moth is more abundant in suburban areas than elsewhere. Overall, our results demonstrate that gardens provide a greater density and diversity of floral resources than farmland, and probably support larger populations of B. terrestris. Contrary to expectation, schemes deployed to enhance farmland biodiversity appear to have little measurable impact on nest growth of this bumblebee species. We argue that B. terrestris probably forage over a larger scale than that on which farms are managed, so that nest growth is determined by the management of a large number of neighbouring farms, not just that in which the nest is located.

In Kinabalu National Park, Borneo we observed four colonies of the Malaysian giant ant Camponotus gigas in a primary forest. These predominantly nocturnal ants have underground nests, but forage in huge three-dimensional territories in the rain forest canopies. The colony on which our study was mainly focused had 17 nests with about 7000 foragers and occupied a territory of 0.8 ha. To improve observation and manipulation possibilities, these nests were linked at ground level by 430 m of artificial bamboo trail. A group of specialist transport worker ants carried food from `source' nests at the periphery to the central `sink' nest of the queen. Transport of food between nests started immediately after the evening exodus of the foragers. Transporter ants formed a physical subcaste among the minors and behaved according to predictions of the central-place foraging theory. Their load size was about five times that of the average forager and grew proportionally with head width. Longer distances were run by ants with greater head width and larger gross weight. Transporter ants that ran more often took heavier loads. Experiments with extra-large baits revealed that C. gigas used long-range recruitment to bring foragers from different nests to "bonanzas" at far distant places. The foraging strategy of C. gigas is based on a polydomous colony structure in combination with efficient communication, ergonomic optimization, polyethism and an effective recruitment system.

Mobile organisms frequently forage for patchy resources; e.g. herbivorous insects searching for host plants. The resource diffusion hypothesis predicts that insect herbivores, such as Pieris rapae butterflies, are disproportionally attracted to more isolated, or ‘diffused’, host plants. Surprisingly little is known about how this response to variation in resource density manifests itself at different spatial scales. We measured the outcome of oviposition by P. rapae butterflies foraging among groups of host plants, with plant density experimentally varied to achieve comparability between three nested scales: fine (1 × 1 m), medium (6 × 6 m), and coarse (36 × 36 m). Hierarchical linear models were used to measure density-dependent responses in the number of eggs laid per plant, with plant density measured at nested spatial scales. At a fine scale, isolated plants received significantly more eggs, while at medium and coarse scales the differences were less pronounced, and tended towards a neutral distribution of eggs across plants. Larger plants also tended to receive more eggs. Since multiple processes, acting at multiple scales, are likely to be the rule rather than the exception in ecology, methods for detecting and characterising multi-scale responses are important to ensure a robust transfer of ecological models from one situation to another.

Competition for food resources can result in spatial and dietary segregation among individuals from the same species. Few studies have looked at such segregations with the combined effect of sex and age in species with short foraging ranges. In this study we examined the 3D spatial use of the environment in a species with a limited foraging area. We equipped 26 little penguins (Eudyptula minor) of known age, sex, and breeding output with GPS (location) and accelerometer (body acceleration and dive depth) loggers. We obtained dietary niche information from the isotopic analysis of blood tissue. We controlled for confounding factors of foraging trip length and food availability by sampling adults at guard stage when parents usually make one-day trips. We observed a spatial segregation between old (>11 years old) and middle-aged penguins (between 5 and 11 years old) in the foraging area. Old penguins foraged closer to the shore, in shallower water. Despite observing age-specific spatial segregation, we found no differences in the diving effort and foraging efficiency between age classes and sexes. Birds appeared to target similar prey types, but showed age-specific variation in their isotopic niche width. We hypothesize that this age-specific segregation was primarily determined by a “cohort effect” that would lead individuals sharing a common life history (i.e. having fledged and dispersed around the same age) to forage preferentially together or to share similar foraging limitations.

Wintering birds can gain significant thermal benefits by foraging in direct sunlight. However, exposure to bright sunlight might make birds easier to detect by predators and may also cause visual glare that can reduce a bird’s ability to monitor the environment. Thus, birds likely experience a trade-off between the thermal benefits and predation-related costs of foraging in direct sunlight. To examine this possible thermoregulation-predation trade-off, we monitored the behavior of mixed-species flocks of wintering emberizid sparrows foraging in alternating strips of sunlight and shade. On average, these sparrows routinely preferred to forage in the shade, despite midday air temperatures as much as 30 °C below their thermoneutral zone. This preference for shade was strongest at relatively high temperatures when the thermal benefits of foraging in sunlight were reduced, suggesting a thermoregulation-predation trade-off. Glare could be reduced if birds faced away from the sun while feeding in direct sunlight, but we found that foraging birds tended to face southward (the direction of the sun). We speculate that other factors, such as the likely direction of predator approach, may explain this southerly orientation, particularly if predators use solar glare to their advantage during an attack. This interpretation is supported by the fact that birds had the weakest southerly orientation on cloudy days. Wintering birds may generally avoid foraging in direct sunlight to minimize their risk of predation. However, given the thermal benefits of sunshine, such birds may benefit from foraging in habitats that provide a mosaic of sunlit and shaded microhabitats.

Invasive plants may compete with native species for abiotic factors as light, space and nutrients, and have also been shown to affect native pollination interactions. Studies have mainly focused on how invasive plants affect pollinator behaviour, i.e. attraction of pollinators to or away from native flowers. However, when an invasive plant provides resources utilized by native pollinators this could increase pollinator population sizes and thereby pollination success in natives. Effects mediated through changes in pollinator population sizes have been largely ignored in previous studies, and the dominance of negative interactions suggested by meta-analyses may therefore be biased. We investigated the impact of the invasive Lupinus polyphyllus on pollination in the native Lotus corniculatus using a study design comparing invaded and uninvaded sites before and after the flowering period of the invasive. We monitored wild bee abundance in transects, and visit rate and seed production of potted Lotus plants. Bumblebee abundance increased 3.9 times in invaded sites during the study period, whereas it was unaltered in uninvaded sites. Total visit rate per Lotus plant increased 2.1 times in invaded sites and decreased 4.4 times in uninvaded sites. No corresponding change in seed production of Lotus was found. The increase in visit rate to Lotus was driven by an increase in solitary bee visitation, whereas mainly bumblebees were observed to visit the invasive Lupinus. The mechanism by which the invasive increases pollinator visit rates to Lotus could be increased availability of other flower resources for solitary bees when bumblebees forage on Lupinus.

Eleven years of data on a small population of adders (Vipera berus) in southern Sweden provide quantitative information on the nature and degree of costs faced by reproducing animals. Reproduction imposes both an energy cost (measured by loss in body mass) and a mortality cost on adders of both sexes. The extent of the energy cost is broadly independent of levels of reproductive activity in males, but mortality costs are highest for large males, perhaps because they are more obvious to predators. In females, energy costs include a high 'fixed' (fecundity-independent) component, such that a large litter may cost little more to produce than would a small litter. Energy costs and mortality costs are separate in males, but inter-related in females. Mortality of reproducing females is high (40% per year), primarily because post-parturient females are emaciated and must forage actively, hence increasing their vulnerability to predators. Females producing relatively large litters (high Relative Clutch Mass) lose more body mass, and are less likely to survive after reproducing. The observed low reproductive frequencies of female adders may result from the presence of high fecundity-independent costs of reproduction.

Parasitoids that forage for herbivorous hosts by using infochemicals may have a problem concerning the reliability and detectability of these stimuli: host stimuli are highly reliable but not very detectable at a distance, while stimuli from the host's food are very detectable but generally not very reliable in indicating host presence. One solution to this problem is to learn to link highly detectable stimuli to reliable but not very detectable stimuli. Ample knowledge is available on how associative learning aids foraging parasitoids in the location of suitable microhabitats. However, in this paper we report on another solution to the reliability-detectability problem and present evidence for an essential, but as yet overlooked, aspect of Drosophila parasitoid ecology. For the first time it is shown that a parasitoid of Drosophila larvae spies on the communication system of adult Drosophila flies to locate potential host sites: naive parasitoids strongly respond to a volatile aggregation pheromone that is deposited in the oviposition site by recently mated female flies. Thus, the parasitoids resort to using highly detectable information from a host stage different from the one under attack (i.e. infochemical detour). The function and ecological implications of these findings are discussed.

In winter, red deer (Cervus elaphus hippelaphus) that are not receiving supplementary feeding emphasize and rely on energy-saving strategies. In severe winters with deep snow, they select a home-range habitat-windblown alpine pastures above the timberline-that meets the requirements of this strategy: a much lower sinking depth and good forage availability are due to patchy snow distribution, while the open area provides higher radiation, permits group formation and thus sociobiological wellbeing. The energy costs of thermoregulation are minimized by selecting slopes with south to south-west exposures, seeking thermal shelter from strong winds, while high flight distances and horizontal flight routes reduce the energy costs of fleeing. In years with little winter snowfall non-supplemented red deer prefer the greater security of forested habitats in lower areas and obtain higher food intake from slightly more nutritious meadows in the valley bottom. The food-supplemented herd has much more constant and predictable habitat use, combining an energy-saving strategy-choosing alpine pastures as low-cost winter home range-with high energy income at the feeding station.

Partially migratory populations, where one portion of a population conducts seasonal migrations (migrants) while the other remains on a single range (residents), are common in ungulates. Studies that assess trade-offs between migratory strategies typically compare the amount of predation risk and forage resources migrants and residents are exposed to only while on separate ranges and assume both groups intermix completely while on sympatric ranges. Here we provide one of the first tests of this assumption by comparing the amount of overlap between home ranges of GPS-collared migrant and resident elk and fine-scale exposure to wolf predation risk and forage biomass at telemetry locations on a sympatric winter range in west-central Alberta, Canada. Overlap between migrant and resident home ranges increased throughout the winter, and both groups were generally intermixed and exposed to equal forage biomass. During the day, both migrants and residents avoided predation risk by remaining in areas far from timber with high human activity, which wolves avoided. However, at night wolves moved onto the grasslands close to humans and away from timber. Resident elk were consistently closer to areas of human activity and further from timber than migrants, possibly because of a habituation to humans. As a result, resident elk were exposed to higher night-time predation risk than migrants. Our study does not support the assumption that migrant and resident elk are exposed to equal predation risk on their sympatric range when human presence alters predation risk dynamics and habituation to humans is unequal between migratory strategies.

Pollinators, even floral generalists (=polyleges), typically specialize during individual foraging bouts, infrequently switching between floral hosts. Such transient floral constancy restricts pollen flow, and thereby gene flow, to conspecific flowers in mixed plant communities. Where incipient flowering species meet, however, weak cross-fertility and often similar floral traits can yield mixed reproductive outcomes among pollinator-dependent species. In these cases, floral constancy by polyleges sometimes serves as an ethological mating barrier. More often, their foraging infidelities instead facilitate host introgression and hybridization. Many other bee species are oligolectic (taxonomic specialists for pollen). Oligoleges could be more discriminating connoisseurs than polyleges when foraging among their limited set of related floral hosts. If true, greater foraging constancy might ensue, contributing to positive assortative mating and disruptive selection, thereby facilitating speciation among their interfertile floral hosts. To test this Connoisseur Hypothesis, nesting females of two species of oligolectic Osmia bees were presented with randomized mixed arrays of flowers of two sympatric species of their pollen host, Balsamorhiza, a genus known for hybridization. In a closely spaced grid, the females of both species preferred the larger flowered B. macrophylla, evidence for discrimination. However, both species’ females showed no floral constancy whatsoever during their individual foraging bouts, switching randomly between species proportional to their floral preference. In a wider spaced array in which the bouquets reflected natural plant spacing, foraging oligolectic bees often transferred pollen surrogates (fluorescent powders) both between conspecific flowers (geitonogamy and xenogamy) and between the two Balsamorhiza species. The Connoisseur Hypothesis was therefore rejected. Foraging infidelity by these oligolectic Osmia bees will contribute to introgression and hybridization where interfertile species of Balsamorhiza meet and flower together. A literature review reveals that other plant genera whose species hybridize also attract numerous oligolectic bees, providing independent opportunities to test the generality of this conclusion.

In the Namib Desert dunes, the web of Seothyra sp. (Eresidae) comprises sticky silk lining the edges of a horizontal mat on the sand surface. The spider sits in a silk-lined burrow attached to the mat. Arthropods become entangled in the sticky silk of the mat and are attacked and pulled into the burrow by the spider. We investigated the influence of sand surface temperature on the activity of spiders during the summer. We determined the range of thermal conditions encountered by spiders, their temperature tolerance and the influence of temperature on foraging activity and prey handling behavior. The environmental temperatures available to Seothyra vary from 17-33° C at the coolest time of day to 33-73° C at the hottest. When prevented from retreating into burrows, spiders showed signs of thermal stress at about 49° C, whereas unrestrained spiders continued to forage at web temperatures above 65° C by moving between the hot surface mat and the cooler burrow. Spiders responded quicker to prey stimuli during the hot hours of the day and completed prey capture sequences in significantly less time at surface temperatures above 49° C than below. Furthermore, captured arthropods succumbed more quickly at high surface temperatures. Our study supports the hypothesis that web design and thermoregulatory behaviors enable Seothyra to hunt under extreme thermal conditions.

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.

Elephants (Loxodonta africana) exhibit pronounced sexual dimorphism, and in this study we test the prediction that the differences in body size and sociality are significant enough to drive divergent foraging strategies and ultimately sexual segregation. Body size influences the foraging behaviour of herbivores through the differential scaling coefficients of metabolism and gut size, with larger bodied individuals being able to tolerate greater quantities of low-quality, fibrous vegetation, whilst having lower mass-specific energy requirements. We test two distinct theories: the scramble competition hypothesis (SCH) and the forage selection hypothesis (FSH). Comprehensive behavioural data were collected from the Pongola Game Reserve and the Phinda Private Game Reserve in South Africa over a 2.5-year period. The data were analysed using sex as the independent variable. Adult females targeted a wider range of species, adopted a more selective foraging approach and exhibited greater bite rates as predicted by the body size hypothesis and the increased demands of reproductive investment (lactation and pregnancy). Males had longer feeding bouts, displayed significantly more destructive behaviour (31% of observations, 11% for females) and ingested greater quantities of forage during each feeding bout. The independent ranging behaviour of adult males enables them to have longer foraging bouts as they experience fewer social constraints than females. The SCH was rejected as a cause of sexual segregation due to the relative abundance of low quality forage, and the fact that feeding heights were similar for both males and females. However, we conclude that the differences in the foraging strategies of the sexes are sufficient to cause spatial segregation as postulated by the FSH. Sexual dimorphism and the associated behavioural differences have important implications for the management and conservation of elephant and other dimorphic species, with the sexes effectively acting as distinct “ecological species”.

Most hypotheses attempting to explain the evolution of reversed sexual dimorphism (RSD) assume that size-related differences in foraging ability are of prime importance, but the studies on sex-specific differences in foraging behaviour remain scarce. We compare the foraging behaviour of males and females in a seabird species with a RSD by using several miniaturised activity and telemetry loggers. In red-footed boobies males are 5% smaller and 15% lighter than females, but have a longer tail than females. Both sexes spend similar time on the nest while incubating or brooding. When foraging at sea, males and females spend similar time foraging in oceanic waters, forage in similar areas, spend similar proportion of their foraging trip in flight, and feed on similar prey-flying fishes and flying squids-of similar size. However, compared to males, females range farther during incubation (85 km vs. 50 km), and furthermore feed mostly at the extremity of their foraging trip, whereas males actively forage throughout the trip. Males are much more active than females, landing and diving more often. During the study period, males lost mass, whereas females showed no significant changes. These results indicate that males and females of the red-footed boobies differ in several aspects in their foraging behaviour. Although some differences found in the study may be the direct result of the larger size of females, that is, the slightly higher speeds and deeper depths attained by females, others indicate clearly different foraging strategies between the sexes. The smaller size and longer tail of males confer them a higher agility, and could allow them to occupy a foraging niche different from that of females. The higher foraging effort of males related to its different foraging strategy is probably at the origin of the rapid mass loss of males during the breeding period. These results suggest that foraging differences are probably the reason for the differential breeding investment observed in boobies, and are likely to be involved in the evolution and maintenance of RSD.

The evolution of queens that rear their first brood solely using body reserves, i.e. fully claustral, is viewed as a major advance for higher ants because it eliminated the need for queens to leave the nest to forage. In an apparently unusual secondary modification, the seed-harvester ant Pogonomyrmex californicus displays obligate queen foraging, i.e. queens must forage to garner the resources necessary to survive and successfully rear their first brood. I examined the potential benefits of queen foraging by comparing ecological and physiological traits between P. californicus and several congeners in which the queen can rear brood using only body reserves. The primary advantage of foraging appears to lie in providing the queens of P. californicus with the energy to raise significantly more brood than possible by congeners that use only body reserves; the workers reared in the first brood were also heavier in mass than that predicted by their head width. Other correlates of queen foraging in P. californicus relative to tested congeners included a significantly lower total fat content for alate queens, a small queen body size, and a low queen to worker body mass ratio. Queens also forage in several other well-studied species of Pogonomyrmex, suggesting the possibility that queen foraging may be more common than previously thought in higher ants.

The foraging decisions of animals are often influenced by risk of predation and by the renewal of resources. For example, seed-eating gerbils on sand dunes in the Negev Desert of Israel prefer to forage in the bush microhabitat and during darker hours due to risk of predation. Also, daily renewal of seed resource patches and timing of nightly foraging activity in a depleting environment play important roles in species coexistence. We examined how these factors influence the timing of gerbil foraging by quantifying foraging activity in seed resource patches that we experimentally renewed hourly during the night. As in previous work, gerbils showed strong preference for the safe bush microhabitat and foraged less in response to high levels of illumination from natural moon light and from artificial sources. We demonstrate here for the first time that gerbils also responded to temporal and spatial heterogeneity in predatory risk through their timing of activity over the course of each night. Typically, gerbils concentrated their activity early in the night, but this changed with moon phase and in response to added illumination. These results can be understood in terms of the nature of patch exploitation by gerbils and the role played by the marginal value of energy in determining the cost of predation. They further show the dynamic nature of gerbil foraging decisions, with animals altering foraging efforts in response to time, microhabital, moon phase, illumination, and resource availability.

We examined whether individual cattle egrets (Bubulcus ibis) base their decisions of where to forage, and how long to stay in a patch, on the behavior of other flock members. Cattle egrets commonly forage in flocks associated with cattle and capture prey at higher rates when they do not share a cow with another egret. Foraging egrets provide cues of the location of prey and their success in capturing prey. Therefore, there is the possibility of information transfer between egrets in a flock. We predicted that egrets should only move to occupied patches when the resident was capturing enough prey that it is profitable for the invader to share the patch or take over the patch. However, egrets did not seem to decide where to forage based on neighbors' rates of energy intake, but rather on the presence or absence of conspecifics in a patch. We also predicted that an egret should remain in a patch until its rate of energy intake dropped to or below the average rate for other egrets within the flock. However, egrets that were foraging more efficiently than the average rate for the flock switched patches sooner than less efficient foragers. Egrets did not appear to increase foraging success by gaining information on patch quality from neighbors.

Predation by naticid gastropods shows evidence of adaptation to maximize the rate of energy intake. The predation rate of Polinices duplicatus feeding on artificially altered, thin-shelled Mercenaria mercenaria was faster than the predation rate on normal Mercenaria. The rate of energy intake was limited by handling time. The time saved by predation on thin-shelled prey was used to forage. Thus time was shown to be valuable to P. duplicatus, and cost-benefit functions using time and energy as currencies are appropriate for estimating dietary efficiency and predicting prey choice.Despite the clear superiority of thin-shelled prey, P. duplicatus did not learn to prefer this novel prey type, suggesting that predator choices are sterotyped, reflecting optima selected over evolutionary time.

Workers of Monomorium minimum forage above-ground for dead arthropods. Small particles (

The rates at which bumblebees of different proboscis lengths forage on flowers of a series of corolla tube lengths were determined. The results indicate significant correlations between proboscis length and time spent by bees on flowers. Bumblebees of long proboscis length can forage significantly faster than bees of shorter proboscis length on flowers with long corolla tubes. There is also evidence which suggests that bumblebees of short proboscis length prefer and are more efficient on short corolla tubes. These results support the use of proboscis length as a morphological indicator of resource utilization in bumblebees.

Two bumblebee species, Bombus bifarius and B. flavifrons, forage randomly with respect to direction when gathering pollen on Potentilla gracilis. Bees avoid revisiting flowers by being able to differentiate recently visited from unvisited flowers. This recognition occurs while bees are flying over open flowers and appears to be a response to the amount of available pollen within flowers. Random foraging with respect to direction is the optimal strategy when the probability of flower revisitation is low. Bumblebees appear to be moving preferentially between nearest neighbors, again as predicted by foraging theory. This behavior causes the establishment of pollen patches in the P. gracilis population. Unlike other pollinators studied in similar situations, bumblebees on P. gracilis do not forage utilizing an area-restricted searching behavior. Because floral reward quality can be assessed at low cost by bees foraging on P. gracilis, their tendency to move to nearby flowers even after encountering a poor quality blossom apparently yields a higher rate of net energy intake than does area-restricted searching. The data indicate that bumblebees exhibit great plasticity in foraging behavior and that they are able to forage efficiently under a wide range of environmental conditions.

The degree of association between cattle egrets (Bubulcus ibis) and cattle was studied during one summer on Saint Catherines Island, Georgia, USA. Previous work by Grubb (1976) and others indicated that cattle egrets foraging with cattle require fewer steps and less time to catch prey than egrets foraging without cattle and single egrets catch prey at a higher rate than egrets foraging in groups of two or more with cattle. Accordingly, we predicted that when given a choice egrets should forage with cattle rather than alone, egrets should prefer to associate with standing rather than sitting cattle, and single egrets associated with cattle should be more common than expected by chance.In excess of two-thirds of the egrets accompanied cattle. Neither time of day nor month influenced the degree of association, but egrets in forest were more likely to be associated with cattle than egrets in pasture. Standing cattle were more likely to be accompanied by egrets than were sitting cattle. Single egrets occurred more frequently than expected by chance when accompanying standing cattle but not when associated with sitting cattle. Thus, cattle egrets usually distributed themselves among cattle in the way predicted by optimal foraging theory.

Four colonies of the desert leaf-cutter ant Acromyrmex versicolor versicolor (Pergande) located 30 miles N.E. of Tempe, Arizona were observed over a 7 month period. The ants utilized trails in foraging, a characteristic of higher attines, as well as foraging singly, a typical pattern among the more primitive gardeners. The ants cut and collected both dry and green vegetation with dry grasses comprising the bulk of the forage. The ants increased their cutting of green vegetation after significant rainfall but collected dry grasses almost exclusively during dry periods. Detailed macro-motion picture analysis of leaf-cutting revealed that the desert gardener utilized a unique technique for cutting compound desert leaves.

Bumblebees foraging on vertical inflorescences start near the bottom and work upward, behavior commonly interpreted as a response to the greater amounts of nectar available in lower flowers. Lupinus polyphyllus, which produces no nectar, has more pollen available in upper flowers. Although bees are probably unable to detect this gradient, since pollen is hidden from their view, they still start low and forage upward. Therefore, we concluded that the bees' tendency to forage upward on vertical inflorescences is not tied to a reward gradient. In addition, bees use only about 15% of the flowers per inflorescence, although they could be much more efficient by visiting and revisiting every flower systematically. In general, revisits would not be penalized because most flowers contain enough pollen for several visits. Optimal foraging theory may not offer an adequate explanation for such gross inefficiency.

The commonest epigeal termite mounds at Nylsvley Reserve, northern Transvaal, are constructed by Cubitermes pretorianus and shared by other termites and by ants. Two surveyed areas had the equivalents of 385 and 496 mounds ha-1 (328 and 432 ha-1 respectively with live Cubitermes). Counts of all termite species and stages in 13 mounds are given. Nests contained up to 15,000 Cubitermes workers (of which up to 0.7% were parasitised by nematodes). Egg-laying was seasonal, with a peak early in the wet season and few eggs in the nest in late autumn (and hence few larvae in winter). Soldier production showed two peaks per year, one at swarming and one in mid-dry season. Aganotermes workers sharing the mounds showed seasonal (over winter) abdominal storage of food reserves. Counts of populations and stored forage in two small Trinervitermes rhodesiensis mounds are also given.