Division of labour and specification of castes in the red imported fire ant Solenopsis invicta buren

Division of labour in Solenopsis invicta follows a familiar pattern: younger, smaller ants tend toward brood care while older, larger ants tend toward foraging. However, long-term observations of marked individuals reveal that length of nursing and foraging ‘careers’ and the age of transition between these activities vary considerably between and within size groups, and are related to length of life. Experiments with entire colonies show that larger ants are more likely than smaller ants to forage for insect prey. There are two main worker castes, ‘nurses’ and ‘foragers’, whose members span a wide age-size range, and a large ‘reserve’ subcaste, heterogeneous in age, size, and behaviour: reserves may nurse, forage, store liquid food, or relay food from nurses to foragers. The proportion of ants engaged in foraging decreases with colony size because many ants in large colonies are not exposed to recruitment signals.     

Dynamics of Social Behavior in Fruit Fly Larvae

We quantified the extent and dynamics of social interactions among fruit fly larvae over time. Both a wild-type laboratory population and a recently-caught strain of larvae spontaneously formed social foraging groups. Levels of aggregation initially increased during larval development and then declined with the wandering stage before pupation. We show that larvae aggregated more on hard than soft food, and more at sites where we had previously broken the surface of the food. Groups of larvae initiated burrowing sooner than solitary individuals, indicating that one potential benefit of larval aggregations is an improved ability to dig and burrow into the food substrate. We also show that two closely related species, D. melanogaster and D. simulans, differ in their tendency to aggregate, which may reflect different evolutionary histories. Our protocol for quantifying social behavior in larvae uncovered robust social aggregations in this simple model, which is highly amenable to neurogenetic analyses, and can serve for future research into the mechanisms and evolution of social behavior.     

Foraging errors play a role in resource exploration by bumble bees (Bombus terrrestris)

If the cognitive performance of animals reflects their particular ecological requirements, how can we explain appreciable variation in learning ability amongst closely related individuals (e.g. foraging workers within a bumble bee colony)? One possibility is that apparent ‘errors’ in a learning task actually represent an alternative foraging strategy. In this study we investigate the potential relationship between foraging ‘errors’ and foraging success among bumble bee (Bombus terrestris) workers. Individual foragers were trained to choose yellow, rewarded flowers and ignore blue, unrewarded flowers. We recorded the number of errors (visits to unrewarded flowers) each bee made during training, then tested them to determine how quickly they discovered a more profitable food source (either familiar blue flowers, or novel green flowers). We found that error prone bees discovered the novel food source significantly faster than accurate bees. Furthermore, we demonstrate that the time taken to discover the novel, more profitable, food source is positively correlated with foraging success. These results suggest that foraging errors are part of an ‘exploration’ foraging strategy, which could be advantageous in changeable foraging environments. This could explain the observed variation in learning performance amongst foragers within social insect colonies.     

Predation risk influences food‐web structure by constraining species diet choice

The foraging behaviour of species determines their diet and, therefore, also emergent food‐web structure. Optimal foraging theory (OFT) has previously been applied to understand the emergence of food‐web structure through a consumer‐centric consideration of diet choice. However, the resource‐centric viewpoint, where species adjust their behaviour to reduce the risk of predation, has not been considered. We develop a mechanistic model that merges metabolic theory with OFT to incorporate the effect of predation risk on diet choice to assemble food webs. This ‘predation‐risk‐compromise’ (PR) model better captures the nestedness and modularity of empirical food webs relative to the classical optimal foraging model. Specifically, compared with optimal foraging alone, risk‐mitigated foraging leads to more‐nested but less‐modular webs by broadening the diet of consumers at intermediate trophic levels. Thus, predation risk significantly affects food‐web structure by constraining species’ ability to forage optimally, and needs to be considered in future work.     

Identification of a gene, Desiccate, contributing to desiccation resistance in Drosophila melanogaster

Suitable alterations in gene expression are believed to allow animals to survive drastic changes in environmental conditions. Drosophila melanogaster larvae cease eating and exit moist food to search for dry pupation sites after the foraging stage in what is known as the wandering stage. Although the behavioral change from foraging to wandering causes desiccation stress, the mechanism by which Drosophila larvae protect themselves from desiccation remains obscure. Here, we identified a gene, CG14686 (designated as Desiccate (Desi)), whose expression was elevated during the wandering stage. The Desi expression level was reversibly decreased by transferring wandering larvae to wet conditions and increased again by transferring them to dry conditions. Elevation of Desi expression was also observed in foraging larvae when they were placed in dry conditions. Desi encoded a 261-amino acid single-pass transmembrane protein with notable motifs, such as SH2 and PDZ domain-binding motifs and a cAMP-dependent protein kinase phosphorylation motif, in the cytoplasmic region, and its expression was observed mainly in the epidermal cells of the larval integuments. Overexpression of Desi slightly increased the larval resistance to desiccation stress during the second instar. Furthermore, Desi RNAi larvae lost more weight under dry conditions, and subsequently, their mortalities significantly increased compared with control larvae. Under dry conditions, consumption of carbohydrate was much higher in Desi RNAi larvae than control larvae. Based on these results, it is reasonable to conclude that Desi contributes to the resistance of Drosophila larvae to desiccation stress.     

Locomotor capacity and foraging behaviour of kalahari lacertid lizards

Closely related lacertid lizards (Eremias, Nucras) in the Kalahari desert differ in patterns of foraging behaviour. Some species are relatively sedentary (‘sit-and-wait’) whereas others are more active (‘widely-foraging’) predators. We determined whether whole-animal locomotor capacities (cruising endurance on a treadmill, initial speed and maximum burst speed in a racetrack, and sprint endurance in a torus-shaped track) correlated with interspecific differences in foraging behaviour. Two of three widely-foraging species had greater cruising endurance, graater sprint endurance, but lower burst speed than did a sit-and-wait species. However, the two species that sprinted quickly also had limited endurance, and vice versa. Pre-feeding negatively influenced endurance but not sprint capacity. Theoretical models of foraging behaviour should recognize that ectotherms have limited endurance, that there can be a trade-off between speed and endurance, and that pre-feeding can reduce some aspects of locomotor capacity.     

Evolution of foraging behaviour in response to chronic malnutrition in Drosophila melanogaster

Chronic exposure to food of low quality may exert conflicting selection pressures on foraging behaviour. On the one hand, more active search behaviour may allow the animal to find patches with slightly better, or more, food; on the other hand, such active foraging is energetically costly, and thus may be opposed by selection for energetic efficiency. Here, we test these alternative hypotheses in Drosophila larvae. We show that populations which experimentally evolved improved tolerance to larval chronic malnutrition have shorter foraging path length than unselected control populations. A behavioural polymorphism in foraging path length (the rover-sitter polymorphism) exists in nature and is attributed to the foraging locus (for). We show that a sitter strain (for(s2)) survives better on the poor food than the rover strain (for(R)), confirming that the sitter foraging strategy is advantageous under malnutrition. Larvae of the selected and control populations did not differ in global for expression. However, a quantitative complementation test suggests that the for locus may have contributed to the adaptation to poor food in one of the selected populations, either through a change in for allele frequencies, or by interacting epistatically with alleles at other loci. Irrespective of its genetic basis, our results provide two independent lines of evidence that sitter-like foraging behaviour is favoured under chronic larval malnutrition.     

Feeding ecology and foraging behaviour of impala Aepyceros melampus in Lake Mburo National Park, Uganda

Intersexual and seasonal variation in foraging behaviour of impala (Aepyceros melampus), was studied in the Lake Mburo National Park, Uganda. There was a moderate seasonal difference in foraging efficiency (as measured by ‘acceptable food abundance’), with a minimum in dry season and a maximum in Rainy season. The variation between sexes was more distinct with a pronounced minimum in time spent browsing of males in early wet season. By distinguishing between feeding time spent grazing and feeding time spent browsing the seasonal variation was confirmed. The proportion of foraging time spent feeding (expressed as ‘food ingestion rate’) showed an inverse pattern with a maximum in the late dry season (75.5%), decreasing values throughout the Rainy season and a minimum in early dry season (57.8%). Differences between sexes were explained in terms of reproductive demands and seasonal balance in terms of moderate climate throughout the year. Impala foraging patterns in the bimodal tropics (two Rainy seasons) is discussed and compared with unimodal tropics. The findings are matched against current ideas on optimal foraging.     

Foraging behaviour in Drosophila larvae: mushroom body ablation

Drosophila larvae and adults exhibit a naturally occurring genetically based behavioural polymorphism in locomotor activity while foraging. Larvae of the rover morph exhibit longer foraging trails than sitters and forage between food patches, while sitters have shorter foraging trails and forage within patches. This behaviour is influenced by levels of cGMP-dependent protein kinase (PGK) encoded by the foraging (for) gene. Rover larvae have higher expression levels and higher PGK activities than do sitters. Here we discuss the importance of the for gene for studies of the mechanistic and evolutionary significance of individual differences in behaviour. We also show how structure-function analysis can be used to investigate a role for mushroom bodies in larval behaviour both in the presence and in the absence of food. Hydroxyurea fed to newly hatched larvae prevents the development of all post-embryonically derived mushroom body (MB) neuropil. This method was used to ablate MBs in rover and sitter genetic variants of foraging to test whether these structures mediate expression of the foraging behavioural polymorphism. We found that locomotor activity levels during foraging of both the rover and sitter larval morphs were not significantly influenced by MB ablation. Alternative hypotheses that may explain how variation in foraging behaviour is generated are discussed.     

Social Learning of a Novel Foraging Skill by White-throated Magpie-jays (Calocitta formosa,Corvidae): a Field Experiment

The cooperatively breeding white-throated magpie-jay (Calocitta formosa) uses a variety of foraging tactics to find, harvest, and process food. Members of territorial groups forage together and may gain information about how to acquire food by observing each other. A field experiment was performed to determine whether a novel skill, door opening to gain access to food, was more rapidly acquired by members of groups in which a trained individual performed the skill. A higher proportion of jays in groups with behavioural ‘models’ (trained birds that opened doors in the presence of group members) acquired the door-opening skill than those groups without models. Young birds acquired the behaviour more frequently than older individuals. Aggressive behaviour at feeders may have affected the spread of the behaviour by reducing the likelihood that individuals performed the behaviour in the presence of other group members but may also have encouraged subordinate individuals to attempt door opening rather than ‘scrounge’.     

Optimal foraging in a uniform habitat: The search mechanism of the green lacewing

The effects of food deprivation and prey contact on the components of searching behaviour in larval green lacewings (Chrysopa carnea Stephens) were examined to test the applicability of optimal foraging theory to predation in a uniform habitat. Variation in foraging intensity was primarily the result of changes in the meander. Modulation of the response to prey contact with increasing deprivation involved changes in the velocity and the response persistence and suggested the occurrence of adaptation to inferred differences in the spatial distribution of the prey. The ratio of giving-up times at different levels of deprivation was in accordance with the predictions of optimal foraging theory, even though the conditions precluded a discrete decision process.     

Evolution of larval foraging behaviour in Drosophila and its effects on growth and metabolic rates

Abstract.  The evolution of foraging in Drosophila melanogaster (Meigen) is studied using outbred populations that had been differentiated using laboratory selection. The foraging behaviour of Drosophila larvae is measured using the foraging path length of 72-h-old larvae. The foraging path length is the distance travelled by foraging larvae over 5 min. Populations of Drosophila selected for rapid development show significantly greater path lengths than their controls. Populations of Drosophila selected for resistance to ammonia and urea in their larval food have shorter path lengths than their controls. Individuals in the ammonia-resistant populations are smaller than those in the control populations, but the size-adjusted metabolic rates are not significantly different. A simple model is proposed suggesting that changes in larval foraging behaviour may be a means for Drosophila larvae to adapt to new environments that require additional maintenance energy. In the ammonia-selected populations, crucial tests of these ideas will have to be conducted in environments with ammonia.     

Selection of Feeding Areas by Senegal Kingfishers Halcyon senegalensis

Studies of ‘optimal foraging’ in birds have concentrated on species which search actively. This paper describes the organization of foraging behaviour under field conditions of a tropical ‘sit-and-wait’ predator, based on the choice of trees as vantage points. Preferences shown for different trees did not appear to maximize the birds' rate of capture of prey items, but rates of energy intake were not measured. However, it is possible that selection for ‘optimal’ behaviour in this species is not so strong as in temperate species which search actively.     

Percentage reward in an operant analogue to foraging

Pigeons (Columba livia) responding on an operant, conditioning analogue to foraging could either accept or reject a variable-interval 20-s schedule which always led to food or a variable-interval 5-s schedule which led to food on only a percentage of trials. The probability of accepting the certain alternative increased as the percentage of food trials for the uncertain alternative decreased. The probability of accepting the uncertain alternative increased with the percentage of food trials for this alternative. Subjects receiving all of their food in the experiment (‘closed economy’) and those requiring supplementation (‘open economy’) preferred whichever alternative provided the higher overall mean rate of reinforcement.     

The vibration dance of the honey bee. I. Communication regulating foraging on two time scales

The relationship between the vibration dance and foraging was investigated for the honey bee, Apis mellifera. Foraging-age workers responded to the vibration dance by moving into the area of the hive where waggle dances were concentrated and by increasing their rate of movement throughout the colony. Vibrated non-foraging-age bees did not move into the waggle dance region or exhibit increased movement in the hive. Small peaks of vibration dance activity, which tended to coincide temporally with small peaks of foraging activity, occurred with a similar frequency throughout the year. These small vibration peaks may have adjusted foraging to short-term fluctuations in food availability. In spring and summer all study hives exhibited large, morning peaks of vibration dance activity, which preceded foraging. Since there was a significant, positive slope for the regression of the magnitude of these morning vibration peaks on the mean level of waggle dancing occurring later during the same day, morning vibration activity may have exerted a long-term ‘priming’ influence on foraging behaviour. In fall and winter, compared with spring and summer, morning vibration dance peaks were smaller, less frequent and tended to coincide with, rather than to precede, foraging activity.     

Social Environment Influences Performance in a Cognitive Task in Natural Variants of the Foraging Gene

In Drosophila melanogaster, natural genetic variation in the foraging gene affects the foraging behaviour of larval and adult flies, larval reward learning, adult visual learning, and adult aversive training tasks. Sitters (for ^s) are more sedentary and aggregate within food patches whereas rovers (for^R) have greater movement within and between food patches, suggesting that these natural variants are likely to experience different social environments. We hypothesized that social context would differentially influence rover and sitter behaviour in a cognitive task. We measured adult rover and sitter performance in a classical olfactory training test in groups and alone. All flies were reared in groups, but fly training and testing were done alone and in groups. Sitters trained and tested in a group had significantly higher learning performances compared to sitters trained and tested alone. Rovers performed similarly when trained and tested alone and in a group. In other words, rovers learning ability is independent of group training and testing. This suggests that sitters may be more sensitive to the social context than rovers. These differences in learning performance can be altered by pharmacological manipulations of PKG activity levels, the foraging (for) gene''s gene product. Learning and memory is also affected by the type of social interaction (being in a group of the same strain or in a group of a different strain) in rovers, but not in sitters. These results suggest that for mediates social learning and memory in D. melanogaster.     

Predation cues rather than resource availability promote cryptic behaviour in a habitat-forming sea urchin

It is well known that predators often influence the foraging behaviour of prey through the so-called “fear effect”. However, it is also possible that predators could change prey behaviour indirectly by altering the prey’s food supply through a trophic cascade. The predator–sea urchin–kelp trophic cascade is widely assumed to be driven by the removal of sea urchins by predators, but changes in sea urchin behaviour in response to predators or increased food availability could also play an important role. We tested whether increased crevice occupancy by herbivorous sea urchins in the presence of abundant predatory fishes and lobsters is a response to the increased risk of predation, or an indirect response to higher kelp abundances. Inside two New Zealand marine reserves with abundant predators and kelp, individuals of the sea urchin Evechinus chloroticus were rarer and remained cryptic (i.e. found in crevices) to larger sizes than on adjacent fished coasts where predators and kelp are rare. In a mesocosm experiment, cryptic behaviour was induced by simulated predation (the addition of crushed conspecifics), but the addition of food in the form of drift kelp did not induce cryptic behaviour. These findings demonstrate that the ‘fear’ of predators is more important than food availability in promoting sea urchin cryptic behaviour and suggest that both density- and behaviourally mediated interactions are important in the predator–sea urchin–kelp trophic cascade.     

Behavioural changes are related to moult regulation in the cockroach, Periplaneta americana

Abstract. Larval holometabolous insects show changes in behaviour (e.g. start of wandering and spinning) in specific periods of the moulting cycle in relation to definite ratios of juvenile hormone and moulting hormone (ecdysone). In hemimetabolous insects no such changes in behaviour are known. It should be investigated whether the cockroach Periplaneta americana shows changes in locomotor activity and in food and water consumption in relation to periods of ecdysone production during the last larval stage. Within a mean duration of the last larval stage of 30 days there were two periods of reduced locomotor activity: on day 9 and between days 13 and 17. From days 12–13 food consumption decreased by c . 40% up to the day 18. Water consumption decreased between days 9 and 18 by about 55%. Peaks of ecdysone production appeared after these changes of behaviour in each case. Therefore in larval Periplaneta ecdysone seems not to trigger these behavioural changes.     

Avoidance of intraspecific competition via host modification in a grazing, fruit-eating insect

Insects feeding on plants may induce chemical and physical changes in the host plants. Here, we present evidence of host plant modification following an insect attack that may be associated with a reduction in intraspecific competition for food. We demonstrate that feeding by larvae of the cranberry fruitworm, Acrobasis vaccinii, induces a change in fruit colour (from green to red) of cranberry fruits, Vaccinium oxycoccos, that is associated with a significant increase in the concentration of anthocyanin. Host fruit colour affected larval foraging behaviour and food acceptance: significantly more cranberry fruitworm larvae were attracted to, and accepted, green rather than red fruits. Our experiments suggest that fruit reddening also prevents exploitation by conspecific larvae of other green fruits adjacent to the attacked fruit.     

Postnatal dispersal of wandering albatrosses Diomedea exulans: implications for the conservation of the species

Many large marine vertebrates are today threatened by human activities and it is therefore crucial to obtain information on their distribution and behaviour at sea. In particular little is known about the time necessary for juveniles to acquire the foraging skills of adults. We tracked 13 juvenile wandering albatrosses Diomedea exulans by satellite telemetry during their first year at sea. They covered an average distance of 184,000 km during the first year and restricted their dispersal to the unproductive waters of the subtropical Indian Ocean and Tasman Sea. This region of low wind velocities does not overlap with the foraging areas used by adults. After an innate phase of rapid dispersal with a fixed flight direction, young birds progressively increased their daily flight distances and attained adult flight efficiency within their first six months at sea. The complete overlap of the juveniles’ foraging ranges with major long‐line fisheries in the subtropical waters constitutes a major threat that could jeopardize the long term recovery ability of populations of the endangered wandering albatross in the Indian Ocean.