Nonrandom foraging and resource distributions affect the relationships between host density, contact rates and parasite transmission.

Nonrandom foraging can cause animals to aggregate in resource dense areas, increasing host density, contact rates and pathogen transmission, but when should nonrandom foraging and resource distributions also have density-independent effects? Here, we used a factorial experiment with constant resource and host densities to quantify host contact rates across seven resource distributions. We also used an agent-based model to compare pathogen transmission when host movement was based on random foraging, optimal foraging or something between those states. Nonrandom foraging strongly depressed contact rates and transmission relative to the classic random movement assumptions used in most epidemiological models. Given nonrandom foraging in the agent-based model and experiment, contact rates and transmission increased with resource aggregation and average distance to resource patches due to increased host movement in search of resources. Overall, we describe three density-independent mechanisms by which host behaviour and resource distributions alter contact rate functions and pathogen transmission.

Disentangling genetic, plastic and social learning drivers of sex-specific foraging behaviour in Trinidadian guppies (Poecilia reticulata).

Evolutionary biologists have long been interested in parsing out the roles of genetics, plasticity and their interaction on adaptive trait divergence. Since males and females often have different ecological and reproductive roles, separating how their traits are shaped by interactions between their genes and environment is necessary and important. Here, we disentangle the sex-specific effects of genetic divergence, developmental plasticity, social learning and contextual plasticity on foraging behaviour in Trinidadian guppies (Poecilia reticulata) adapted to high- or low-predation habitats. We reared second-generation siblings from both predation regimes with or without predator chemical cues, and with adult conspecifics from either high- or low-predation habitats. We then quantified their foraging behaviour in water with and without predator chemical cues. We found that high-predation guppies forage more efficiently than low-predation guppies, but this behavioural difference is context-dependent and shaped by different mechanisms in males and females. Higher foraging efficiency in high-predation females is largely genetically determined, and to a smaller extent socially learned from conspecifics. However, in high-predation males, higher foraging efficiency is plastically induced by predator cues during development. Our study demonstrates sex-specific differences in genetic versus plastic responses in foraging behaviour, a trait of significance in organismal fitness and ecosystem dynamics.

High toxin concentration in pollen may deter collection by bees in butterfly-pollinated Rhododendron molle.

BACKGROUNDS AND AIMS: The hypothesis that plants evolve features that protect accessible pollen from consumption by flower visitors remains poorly understood. METHODS: To explore potential chemical defenses against pollen consumption, we examined the pollinator assemblage, foraging behaviour, visitation frequency and pollen transfer efficiency in Rhododendron molle, a highly toxic shrub containing Rhodojaponin III. Nutrient (protein and lipid) and toxic components in pollen and other tissues were measured. KEY RESULTS: Overall in the five populations, floral visits by butterflies and bumblebees were relatively more frequent than visits by honeybees. All foraged for nectar but not pollen. Butterflies did not differ from bumblebees in the amount of pollen removed per visit, but deposited more pollen per visit. Pollination experiments indicated that R. molle was self-compatible, but both fruit and seed production were pollen limited. Our analysis indicated that the pollen was not protein-poor and had a higher concentration of the toxic compound Rhodojaponin III than petals and leaves, which compound was undetectable in nectar. CONCLUSION: Pollen toxicity in Rhododendron flowers may discourage pollen robbers (bees) from taking the freely accessible pollen grains, while the toxin-free nectar rewards effective pollinators, promoting pollen transfer. This preliminary study supports the hypothesis that chemical defense in pollen would be likely to evolve in species without physical protection from pollinivores.

Resource manipulation reveals interactive phenotype-dependent foraging in free-ranging lizards.

Recent evidence suggests that individuals differ in foraging tactics and this variation is often linked to an individual's behavioural type (BT). Yet, while foraging typically comprises a series of search and handling steps, empirical investigations have rarely considered BT-dependent effects across multiple stages of the foraging process, particularly in natural settings. In our long-term sleepy lizard (Tiliqua rugosa) study system, individuals exhibit behavioural consistency in boldness (measured as an individual's willingness to approach a novel food item in the presence of a threat) and aggressiveness (measured as an individual's response to an 'attack' by a conspecific dummy). These BTs are only weakly correlated and have previously been shown to have interactive effects on lizard space use and movement, suggesting that they could also affect lizard foraging performance, particularly in their search behaviour for food. To investigate how lizards' BTs affect their foraging process in the wild, we supplemented food in 123 patches across a 120-ha study site with three food abundance treatments (high, low and no-food controls). Patches were replenished twice a week over the species' entire spring activity season and feeding behaviours were quantified with camera traps at these patches. We tracked lizards using GPS to determine their home range (HR) size and repeatedly assayed their aggressiveness and boldness in designated assays. We hypothesised that bolder lizards would be more efficient foragers while aggressive ones would be less attentive to the quality of foraging patches. We found an interactive BT effect on overall foraging performance. Individuals that were both bold and aggressive ate the highest number of food items from the foraging array. Further dissection of the foraging process showed that aggressive lizards in general ate the fewest food items in part because they visited foraging patches less regularly, and because they discriminated less between high and low-quality patches when revisiting them. Bolder lizards, in contrast, ate more tomatoes because they visited foraging patches more regularly, and ate a higher proportion of the available tomatoes at patches during visits. Our study demonstrates that BTs can interact to affect different search and handling components of the foraging process, leading to within-population variation in foraging success. Given that individual differences in foraging and movement will influence social and ecological interactions, our results highlight the potential role of BT's in shaping individual fitness strategies and population dynamics.

Physiological tipping points in the relationship between foraging success and lifetime fitness of a long-lived mammal.

Although anthropogenic change is often gradual, the impacts on animal populations may be precipitous if physiological processes create tipping points between energy gain, reproduction or survival. We use 25 years of behavioural, diet and demographic data from elephant seals to characterise their relationships with lifetime fitness. Survival and reproduction increased with mass gain during long foraging trips preceding the pupping seasons, and there was a threshold where individuals that gained an additional 4.8% of their body mass (26 kg, from 206 to 232 kg) increased lifetime reproductive success three-fold (from 1.8 to 4.9 pups). This was due to a two-fold increase in pupping probability (30% to 76%) and a 7% increase in reproductive lifespan (6.0 to 6.4 years). The sharp threshold between mass gain and reproduction may explain reproductive failure observed in many species and demonstrates how small, gradual reductions in prey from anthropogenic disturbance could have profound implications for animal populations.

Monarch butterflies memorize the spatial location of a food source.

Spatial memory helps animals to navigate familiar environments. In insects, spatial memory has extensively been studied in central place foragers such as ants and bees. However, if butterflies memorize a spatial location remains unclear. Here, we conducted behavioural experiments to test whether monarch butterflies (Danaus plexippus) can remember and retrieve the spatial location of a food source. We placed several visually identical feeders in a flight cage, with only one feeder providing sucrose solution. Across multiple days, individual butterflies predominantly visited the rewarding feeder. Next, we displaced a salient landmark close to the feeders to test which visual cue the butterflies used to relocate the rewarding feeder. While occasional landmark displacements were ignored by the butterflies and did not affect their decisions, systematic displacement of both the landmark and the rewarding feeder demonstrated that the butterflies associated the salient landmark with the feeder's position. Altogether, we show that butterflies consolidate and retrieve spatial memory in the context of foraging.

Bee pollination services and the burden of biogeography.

Native bees augment pollination services in the Northern Hemisphere, especially cultivated apple crops, yet Southern Hemisphere contexts are poorly known. We observed the foraging behaviour of 69 354 invertebrate flower visitors in Australian orchards (two regions, 3 years) to assess the efficacy of pollination service (Peff). Native stingless bees and introduced honey bees were the most abundant visitors and most efficacious pollinators (Tetragonula Peff = 6.16; Apis Peff = 13.02), with Tetragonula becoming important service providers above 22°C. However, visits by tree-nesting stingless bees decreased with distance from native forest (less than 200 m) and their tropical/subtropical distribution precludes pollination service in other major Australian apple-producing regions. More broadly distributed native allodapine and halictine bees transferred the most pollen per-visit, but their low abundances reduce efficacies (Exoneura Peff = 0.03; Lasioglossum Peff = 0.06), resulting in a general dependence on honey bees. This reliance is a burden of biogeography, since key Northern Hemisphere pollinators of apple (Andrena, Apis, Bombus, Osmia) do not naturally occur in Australasia-where there is only 15% generic overlap with Central Asian bees sympatric with wild apple distributions (cf. Palaearctic 66% and Nearctic 46% generic overlaps). The historical biogeography of bees therefore drives an extreme dependence on one introduced species for apple pollination in Australia.

Acoustic activity of bats at power lines correlates with relative humidity: a potential role for corona discharges.

With the ever-increasing dependency on electric power, electrical grid networks are expanding worldwide. Bats exhibit a wide diversity of foraging and flight behaviours, and their sensitivity to anthropogenic stressors suggests this group is very likely to be affected by power lines in a myriad of ways. Yet the effects of power lines on bats remains unknown. Here we assessed the responses of insectivorous bats to very high voltage power lines (VHVPL; greater than 220 kV). We implemented a paired sampling design and monitored bats acoustically at 25 pairs, one pair consisting of one forest edge near to VHVPL matched with one control forest edge. Relative humidity mediates the effects of power lines on bats: we detected bat attraction to VHVPL at high relative humidity levels and avoidance of VHVPL by bats at low relative humidity levels. We argue that the former could be explained by insect attraction to the light emitted by VHVPL owing to corona discharges while the latter may be owing to the physical presence of pylons/cables at foraging height and/or because of electromagnetic fields. Our work highlights the response of bats to power lines at foraging habitats, providing new insight into the interactions between power lines and biodiversity.

Pollinator foraging tactics have divergent consequences for the mating system of a tropical plant.

Resolving the consequences of pollinator foraging behaviour for plant mating systems is a fundamental challenge in evolutionary ecology. Pollinators may adopt particular foraging tactics: complete trapline foraging (repeated movements along a fixed route), sample-and-shift trapline foraging (a variable route that incorporates information from previous experiences) and territorial foraging (stochastic movements within a restricted area). Studies that integrate these pollinator foraging tactics with plant mating systems are generally lacking. We investigate the consequences of particular pollinator foraging tactics for Heliconia tortuosa. We combine parentage and sibship inference analysis with simulation modelling to: estimate mating system parameters; infer the foraging tactic adopted by the pollinators; and quantify the impact of pollinator foraging tactics on mating system parameters. We found high outcrossing rates, ubiquitous multiple paternity and a pronounced departure from near-neighbour mating. We also found that plants repeatedly receive pollen from a series of particular donors. We infer that the pollinators primarily adopt complete trapline foraging and occasionally engage in sample-and-shift trapline foraging. This enhances multiple paternity without a substantial increase in near-neighbour mating. The particular pollinator foraging tactics have divergent consequences for multiple paternity and near-neighbour mating. Thus, pollinator foraging behaviour is an important driver of the ecology and evolution of plant mating systems.

Omega-3 enriched chick diet reduces the foraging areas of breeders in two closely related shearwaters from contrasting marine environments.

Seabirds have evolved several life-history characteristics to help buffer environmental stochasticity. However, particularly during the breeding season, seabirds may be affected by reductions in prey availability and localised oceanographic conditions caused by variations in the environment. The increase in sea surface temperature, triggered by accelerated global warming, is impairing phytoplankton production of omega-3 fatty acids (FAs). Here, we assessed the ecological role of omega-3 FAs on chick development and subsequently on breeder foraging behaviour in two closely related shearwater species foraging in contrasting marine environments. We supplemented chicks with omega-3 FA pills or with control placebo pills and monitored chick growth, chick health status and breeder at-sea foraging behaviour using global positioning system devices. We found that omega-3 chick supplementation reduced the 95% kernel utilization distribution of short trips of Cape Verde shearwaters, but overall, breeders kept a similar foraging pattern between treatments, potentially influenced by predictable prey patches off the West African coast. In contrast, for Cory's shearwaters, the parents of the omega-3 group greatly reduced the foraging effort. This suggests that the proximity to productive prey patches around the colony may help birds to adjust their effort and, therefore, energy expenditure, to changes in the development of their offspring, as driven by their nutritional status. Overall, our results suggest a link between a chick diet enriched in omega-3 FAs and parental foraging effort, providing insight into their ability to cope with a changing and increasingly stochastic marine environment.

Boldness predicts plasticity in flight responses to winds.

Behavioural plasticity can allow populations to adjust to environmental change when genetic evolution is too slow to keep pace. However, its constraints are not well understood. Personality is known to shape individual behaviour, but its relationship to behavioural plasticity is unclear. We studied the relationship between boldness and behavioural plasticity in response to wind conditions in wandering albatrosses (Diomedea exulans). We fitted multivariate hidden Markov models to an 11-year GPS dataset collected from 294 birds to examine whether the probability of transitioning between behavioural states (rest, prey search and travel) varied in response to wind, boldness and their interaction. We found that movement decisions varied with boldness, with bolder birds showing preferences for travel, and shyer birds showing preferences for search. For females, these effects depended on wind speed. In strong winds, which are optimal for movement, females increased time spent in travel, while in weaker winds, shyer individuals showed a slight preference for search, while bolder individuals maintained preference for travel. Our findings suggest that individual variation in behavioural plasticity may limit the capacity of bolder females to adjust to variable conditions and highlight the important role of behavioural plasticity in population responses to climate change.

Following the feeder: A global synthesis of disturbance-based foraging associations of birds.

Species interactions link animal behaviour to community structure and macroecological patterns of biodiversity. One common type of trophic species interaction is disturbance foraging-the act of obtaining food at a disturbance created by another organism. Disturbance foraging is widespread across the animal kingdom, especially among birds, yet previous research has been largely anecdotal and we still lack a synthetic understanding of how this behaviour varies geographically, phylogenetically and ecologically. To address these gaps, we conducted a comprehensive literature review to test focal hypotheses about disturbance foraging behaviour in birds. We found that avian disturbance foraging was geographically ubiquitous, occurring in both aquatic and terrestrial habitats across six continents and four oceans. Consistent with predictions based on established species diversity gradients in different habitat types, the majority of terrestrial observations occurred at tropical latitudes, whereas aquatic observations took place most frequently in temperate marine waters. Although disturbance foraging was widespread across the avian phylogeny, contrary to our prediction, the behaviour was also conserved phylogenetically (Pagel's λ = 0.7) and clustered within suboscine landbirds in terrestrial environments and seabirds in aquatic environments. Similarly, although disturbers were taxonomically diverse as we predicted, interactions were unexpectedly dominated by swarm-raiding ants in terrestrial environments and cetaceans in aquatic environments. Diet and body mass were also important predictors of disturbance foraging associations: Responders followed disturbers with similar diets and larger body sizes. Overall, our hypothesis-testing framework provides insight into the importance of geography, phylogeny and ecology as predictors of disturbance foraging behaviour. We anticipate that this comprehensive assessment of disturbance foraging will serve to generate additional hypotheses and spark future research and management considerations about this fascinating but poorly studied suite of species interactions, especially as biotic interactions face unprecedented risks in our rapidly changing world.

Exaggerated mandibles are correlated with enhanced foraging efficacy in male Auckland tree weta.

Sexual selection has driven the evolution of weaponry for males to fight rivals to gain access to females. Although weapons are predicted to increase males' reproductive success, they are also expected to incur costs and may impair functional activities, including foraging. Using feeding assays, we tested whether the enlarged mandibles of Auckland tree weta (Hemideina thoracica) impact feeding activity (the total volume of biomass consumed, bite rate, and number of foraging visits) and foraging behaviour (time spent moving, feeding, or stationary). We predicted that increased head capsule size in male weta would hinder their foraging efficacy. However, we found that weta with longer heads fed at a faster rate and spent less time foraging than weta with smaller heads, regardless of sex. Contrary to expectations that weapons impede functional activities, our results demonstrate that exaggerated traits can improve feeding performance and may offer benefits other than increased mating success.

Larger pollen loads increase risk of heat stress in foraging bumblebees.

Global declines in bumblebee populations are linked to climate change, but specific mechanisms imposing thermal stress on these species are poorly known. Here we examine the potential for heat stress in workers foraging for pollen, an essential resource for colony development. Laboratory studies have shown that pollen foraging causes increased thoracic temperatures (Tth) in bees, but this effect has not been examined in bumblebees nor in real-world foraging situations. We examine the effects of increasing pollen load size on Tth of Bombus impatiens workers in the field while accounting for body size and microclimate. We found that Tth increased by 0.07°C for every milligram of pollen carried (p = 0.007), resulting in a 2°C increase across the observed range of pollen load sizes. Bees carrying pollen were predicted to have a Tth 1.7-2.2°C hotter than those without pollen, suggesting that under certain conditions, pollen loads could cause B. impatiens workers to heat from a safe Tth to one within the range of their critical thermal limits that we measured (41.3°C to 48.4°C). Bumblebees likely adopt behavioural or physiological strategies to counteract the thermal stress induced by pollen transport, and these may limit their foraging opportunities as environmental temperatures continue to increase.

Extremely low seasonal prey capture efficiency in a deep-diving whale, the narwhal.

Successful foraging is essential for individuals to maintain the positive energy balance required for survival and reproduction. Yet, prey capture efficiency is poorly documented in marine apex predators, especially deep-diving mammals. We deployed acoustic tags and stomach temperature pills in summer to collect concurrent information on presumed foraging activity (through buzz detection) and successful prey captures (through drops in stomach temperature), providing estimates of feeding efficiency in narwhals. Compared to the daily number of buzzes (707 ± 368), the daily rate of feeding events was particularly low in summer (19.8 ± 8.9) and only 8-14% of the foraging dives were successful (i.e. with a detectable prey capture). This extremely low success rate resulted in a very low daily food consumption rate (less than 0.5% of body mass), suggesting that narwhals rely on body reserves accumulated in winter to sustain year-round activities. The expected changes or disappearance of their wintering habitats in response to climate change may therefore have severe fitness consequences for narwhal populations.

Spatiotemporal variation in the role of floral traits in shaping tropical plant-pollinator interactions.

The pollination syndrome hypothesis predicts that plants pollinated by the same pollinator group bear convergent combinations of specific floral functional traits. Nevertheless, some studies have shown that these combinations predict pollinators with relatively low accuracy. This discrepancy may be caused by changes in the importance of specific floral traits for different pollinator groups and under different environmental conditions. To explore this, we studied pollination systems and floral traits along an elevational gradient on Mount Cameroon during wet and dry seasons. Using Random Forest (Machine Learning) models, allowing the ranking of traits by their relative importance, we demonstrated that some floral traits are more important than others for pollinators. However, the distribution and importance of traits vary under different environmental conditions. Our results imply the need to improve our trait-based understanding of plant-pollinator interactions to better inform the debate surrounding the pollination syndrome hypothesis.

Top-down effects of foraging decisions on local, landscape and regional biodiversity of resources (DivGUD).

Foraging by consumers acts as a biotic filtering mechanism for biodiversity at the trophic level of resources. Variation in foraging behaviour has cascading effects on abundance, diversity, and functional trait composition of the community of resource species. Here we propose diversity at giving-up density (DivGUD), i.e. when foragers quit exploiting a patch, as a novel concept and simple measure quantifying cascading effects at multiple spatial scales. In experimental landscapes with an assemblage of plant seeds, patch residency of wild rodents decreased local α-DivGUD (via elevated mortality of species with large seeds) and regional γ-DivGUD, while dissimilarity among patches in a landscape (ß-DivGUD) increased. By linking theories of adaptive foraging behaviour with community ecology, DivGUD allows to investigate cascading indirect predation effects, e.g. the ecology-of-fear framework, feedbacks between functional trait composition of resource species and consumer communities, and effects of inter-individual differences among foragers on the biodiversity of resource communities.

Elephant seal foraging success is enhanced in Antarctic coastal polynyas.

Antarctic polynyas are persistent open water areas which enable early and large seasonal phytoplankton blooms. This high primary productivity, boosted by iron supply from coastal glaciers, attracts organisms from all trophic levels to form a rich and diverse community. How the ecological benefit of polynya productivity is translated to the highest trophic levels remains poorly resolved. We studied 119 southern elephant seals feeding over the Antarctic shelf and demonstrated that: (i) 96% of seals foraging here used polynyas, with individuals spending on average 62% of their time there; (ii) the seals exhibited more area-restricted search behaviour when in polynyas; and (iii) these seals gained more energy (indicated by increased buoyancy from greater fat stores) when inside polynyas. This higher-quality foraging existed even when ice was not present in the study area, indicating that these are important and predictable foraging grounds year-round. Despite these energetic advantages from using polynyas, not all the seals used them extensively. Factors other than food supply may influence an individual's choice in their use of feeding grounds, such as exposure to predation or the probability of being able to return to distant sub-Antarctic breeding sites.

Scavenging in the realm of senses: smell and vision drive recruitment at carcasses in Neotropical ecosystems.

Social information, acquired through the observation of other individuals, is especially relevant among species belonging to the same guild. The unpredictable and ephemeral nature of carrion implies that social mechanisms may be selected among scavenger species to facilitate carcass location and consumption. Here, we apply a survival-modelling strategy to data obtained through the placement and monitoring of carcasses in the field to analyse possible information transmission cascades within a Neotropical scavenger community. Our study highlights how the use of different senses (smell and sight) within this guild facilitates carcass location through the transmission of social information between species with different carrion foraging efficiencies. Vultures with a highly developed sense of smell play a key role in this process, as they are the first to arrive at the carcasses and their presence seems to serve as a visual cue for other species to locate the resource. Our study supports the local enhancement hypothesis within scavengers, whereby individuals locate carcasses by following foraging heterospecifics, also suggesting the importance of the sense of smell in the maintenance of the community structure.

Memory-guided foraging and landscape design interact to determine ecosystem services.

Many studies examine how the landscape affects memory-informed movement patterns, but very few examine how memory-informed foragers influence the landscape. This reverse relationship is an important factor in preventing the continued decline of many ecosystem services. We investigate this question in the context of crop pollination services by wild bees, a critical ecosystem service that is in steep decline. Many studies suggest that adding wild flower patches near crops can result in higher crop pollination services, but specific advice pertaining to the optimal location and density of these wild flower patches is lacking, as well as any estimate of the expected change in crop pollination services. In this work, we seek to understand what is the optimal placement of a flower patch relative to a single crop field, during crop bloom and considering spatial factors alone. We develop an individual based model of memory-based foraging by bumble bees to simulate bee movement from a single nest while the crop is in bloom, and measure the resulting crop pollination services. We consider a single crop field enhanced with a wild flower patch in a variable location, and measure crop flower visitation over the course of a single day. We analyze the pollination intensity and spatial distribution of flower visits to determine optimal wild flower patch placement for an isolated crop field. We find that the spatial arrangement of crop and wild flower patch have a significant effect on the number of crop flower visits, and that these effects arise from the memory-informed foraging pattern. The most effective planting locations are either in the centre of the crop field or on the far side of the crop field, away from the single bumble bee nest.