Traplining by purple-throated carib hummingbirds: behavioral responses to competition and nectar availability

We examined the effects of nectar availability and competition on foraging preferences and revisit intervals of traplining female purple-throated caribs hummingbirds (Eulampis jugularis) to Heliconia patches shared by two individuals or visited solely by one individual. Birds at both shared and solitary patches preferred multiflowered to single-flowered inflorescences, but the magnitude of this preference depended on food availability and competition. During a year of low flower availability, females visited multiflowered inflorescences more frequently than single-flowered inflorescences only when nectar availability was experimentally enhanced; similarly, females at shared patches exhibited a significant preference for multiflowered inflorescences only after experimental increases in nectar availability. Experimental manipulations of nectar availability also had different effects on revisit intervals of birds at shared vs solitary patches. Birds at shared patches responded to patch-wide increases in nectar rewards by increasing the duration of their visit intervals, whereas birds at solitary patches did not. In contrast, birds at solitary patches responded to abrupt losses of nectar at flowers (simulating competition) by decreasing the duration of their visit intervals, whereas a bird at a shared patch did not alter its return interval. The contrasting results between shared vs solitary patches suggest that future studies of traplining behavior should incorporate levels of competition into their design.     

What makes a honeybee scout?

A honeybee colony needs to divide its workforce so that each of the many tasks it performs has an appropriate number of workers assigned to it. This task allocation system needs to be flexible enough to allow the colony to quickly adapt to an ever-changing environment. In this study, we examined possible mechanisms by which a honeybee colony regulates the division of labor between scouts (foragers that search for new food sources without having been guided to them) and recruits (foragers that were guided via recruitment dances toward food sources). Specifically, we examined the roles that the availability of recruitment dances and worker genotype has in the colony-level regulation of the number of workers engaged in scouting. Our approach was threefold. We first developed a mathematical model to demonstrate that the decision to become a scout or a recruit could be regulated by whether a potential forager can find a recruitment dance within a certain time period. We then tested this model by investigating the effect of dance availability on the regulation of scouts in the field. Lastly, we investigated if the probability of being a scout has a genetic basis. Our field data supported the hypothesis that scouts are those foragers that have failed to locate a recruitment dance as predicted by our model, but we found no effect of genotype on the propensity of foragers to become scouts.     

Why do snakes have eyes? The (non-)effect of blindness in island tiger snakes (Notechis scutatus)

Large (to >1 m), diurnally active tiger snakes (Notechis scutatus) are abundant on Carnac Island, near the coast of Western Australia. Our behavioural and mark-recapture studies provide the first ecological data on this population, and reveal a surprising phenomenon. Many adult tiger snakes have had their eyes destroyed, apparently during nest defence, by silver gulls (Larus novaehollandiae). This loss of vision did not reduce the snakes' body condition (mass relative to length), or their rates of growth or survival (measured over a 12-month period). Blind male snakes trail-followed females, and mated successfully. Thus, destruction of a major sensory modality had no detectable effect on these predators. This result is strongly counter-intuitive, but mirrors an earlier report of congenital blindness (without ill-effects) in American viperid snakes. Similarities between the two systems (island populations, highly venomous snakes, reliance on sessile prey) clarify the circumstances under which the loss of vision does not reduce an organism's viability. These natural experiments support Gans' hypothesis of “momentarily excessive construction” in that the snakes possess a complex organ system that they do not actually require for successful feeding, survival or reproduction. Received: 30 November 1998 / Received in revised form: 24 April 1999 / Accepted: 9 May 1999     

Pattern-oriented modeling of bird foraging and pest control in coffee farms

We develop a model of how land use and habitat diversity affect migratory bird populations and their ability to suppress an insect pest on Jamaican coffee farms. Bird foraging—choosing which habitat patch and prey to use as prey abundance changes over space and time—is the key process driving this system. Following the “pattern-oriented” modeling strategy, we identified nine observed patterns that characterize the real system's dynamics. The model was designed so that these patterns could potentially emerge from it. The resulting model is individual-based, has fine spatial and temporal resolutions, represents very simply the supply of the pest insect and other arthropod food in six habitat types, and includes foraging habitat selection as the only adaptive behavior of birds. Although there is an extensive heritage of bird foraging theory in ecology, most of it addresses only the individual level and is too simple for our context. We used pattern-oriented modeling to develop and test foraging theory for this across-scale problem: rules for individual bird foraging that cause the model to reproduce a variety of patterns observed at the system level. Four alternative foraging theories were contrasted by how well they caused the model to reproduce the nine characteristic patterns. Four of these patterns were clearly reproduced with the “null” theory that birds select habitat randomly. A version of classical theory in which birds stay in a patch until food is depleted to some threshold caused the model to reproduce five patterns; this theory caused lower, not higher, use of habitat experiencing an outbreak of prey insects. Assuming that birds select the nearby patch providing highest intake rate caused the model to reproduce all but one pattern, whereas assuming birds select the highest-intake patch over a large radius produced an unrealistic distribution of movement distances. The pattern reproduced under none of the theories, a negative relation between bird density and distance to trees, appears to result from a process not in the model: birds return to trees at night to roost. We conclude that a foraging model for small insectivorous birds in diverse habitat should assume birds can sense higher food supply but over short, not long, distances.     

Brood pheromone modulates honeybee (Apis mellifera L.) sucrose response thresholds

Foraging behavior and the mechanisms that regulate foraging activity are important components of social organization. Here we test the hypothesis that brood pheromone modulates the sucrose response threshold of bees. Recently the honeybee proboscis extension response to sucrose has been identified as a ”window” into a bee’s perception of sugar. The sucrose response threshold measured in the first week of adult life, prior to foraging age, predicts forage choice. Bees with low response thresholds are more likely to be pollen foragers and bees with high response thresholds are more likely to forage for nectar. There is an associated genetic component to sucrose response thresholds and forage choice such that bees selected to hoard high quantities of pollen have low response thresholds and bees selected to hoard low quantities of pollen have higher response thresholds. The number of larvae in colonies affects the number of bees foraging for pollen. Hexane-extractable compounds from the surface of larvae (brood pheromone) significantly increase the number of pollen foragers. We tested the hypothesis that brood pheromone decreases the sucrose response threshold of bees, to suggest a pheromone- modulated sensory-physiological mechanism for regulating foraging division of labor. Brood pheromone significantly decreased response thresholds as measured in the proboscis extension response assay, a response associated with pollen foraging. A synthetic blend of honeybee brood pheromone stimulated and released pollen foraging in foraging bioassays. Synthetic brood pheromone had dose-dependent effects on the modulation of sucrose response thresholds. We discuss how brood pheromone may act as a releaser of pollen foraging in older bees and a primer pheromone on the development of response thresholds and foraging ontogeny of young bees. Received: 24 May 2000 / Revised: 26 September 2000 / Accepted: 15 October 2000     

Hunger-dependent predator inspection and foraging behaviours in the threespine stickleback (Gasterosteus aculeatus) under predation risk

Individual fish commonly leave the relative safety of the shoal to approach potential predators at a distance. Not all members of a shoal are equally likely to initiate such predator inspection visits. Here, we show for the first time that the current hunger state of individual fish strongly influences their predator inspection behaviour, as well as their foraging rate, in the face of predation hazard. When all members of threespine stickleback (Gasterosteus aculeatus) test shoals were in a similar hunger state, they were equally likely to inspect a trout predator model alone and did not differ in the frequency of their inspection visits or foraging rate. However, when individual sticklebacks in a shoal differed in their hunger state, the food-deprived (i.e. hungrier) member of the shoal fed at a higher rate, was significantly more likely to initiate solitary predator inspection visits, and inspected the predator model significantly more often than its less hungry (i.e. well-fed) shoal mates. Individual fish which inspected the predator model more frequently also tended to have higher feeding rates. The results indicate that the hungrier fish in a shoal are more willing to take greater risks to inspect a potential threat at a distance, compared with their well-fed shoal mates, and suggest that they may gain a foraging benefit in doing so. If marked asymmetries in hunger state exist among members of fish shoals, then mutual cooperation during predator inspection visits may be difficult to achieve because well-fed individuals are not as likely to initiate or participate in inspection visits as are hungry individuals.Correspondence to: J.-G.J. Godin     

Yellowjackets (Vespula pensylvanica) thermoregulate in response to changes in protein concentration

Social insects can modulate body temperature to increase foraging efficiency; however, little is known about how the relative value of protein resources affects forager body temperature. Such regulation may be important given that colony growth is often limited by protein availability. In this paper, we present what are, to our knowledge, the first data for social insects showing that thoracic temperatures (T (th)) of foragers increase with the protein content of food resources. In an introduced population of western yellowjacket (Vespula pensylvanica), we measured T (th) of foragers collecting high-quality protein (100% canned chicken) and low-quality protein (50% canned chicken, 50% indigestible alpha-cellulose by volume) at different ambient air temperatures (T (a)). Wasps foraging on 100% chicken consistently exhibited higher T (th) compared to wasps foraging on 50% chicken. After correcting for T (a), the mean T (th) for wasps collecting 100% chicken were 1.98 degrees C higher than those of individuals collecting 50% chicken. We suggest that this mechanism may increase foraging efficiency in this and other social wasp species.     

Coming out of the woods: do termites need a specialized worker caste to search for new food sources?

Most small-colony termites live confined within a single piece of wood on which they feed and do not possess permanent workers: Tasks are done by developmentally flexible immatures (pseudergates). By contrast, large-colony termites possess a specialized (true) worker caste and forage outside their nest for food. To shed light on possible transitional steps between these contrasting patterns of social organization, we studied an atypical Rhinotermitidae, Prorhinotermes inopinatus. In this species, despite the absence of a true worker caste, soldiers, pseudergates, and neotenic reproductives may leave the nest and explore their surroundings. Although evidence presented in this study indicates that termites recognize unknown areas, there is no directional recruitment toward them. The discovery of a food source, i.e., a piece of wood, is followed by the establishment of a long-lasting trail between the nest and the food source. A large fraction of the colony, including neotenic reproductives, ultimately migrates into the piece of wood. Our results thus demonstrate that multiple features of external foraging behavior can evolve independently of the existence of a true worker caste in termites. We suggest that large colonies with true workers, like those of most Rhinotermitidae, may easily have evolved from a Prorhinotermes-like pattern if submitted to increasing selective pressures for worker efficiency in a stable environment.     

The effects of colony-level selection on the social organization of honey bee (Apis mellifera L.) colonies: colony-level components of pollen hoarding

Two-way selection for quantities of stored pollen resulted in the production of high and low pollen hoarding strains of honey bees (Apis mellifera L.). Strains differed in areas of stored pollen after a single generation of selection and, by the third generation, the high strain colonies stored an average 6 times more pollen than low strain colonies. Colony-level organizational components that potentially affect pollen stores were identified that varied genetically within and between these strains. Changes occurred in several of these components, in addition to changes in the selected trait. High strain colonies had a significantly higher proportion of foragers returning with loads of pollen, however, high and low strain colonies had equal total numbers of foragers Colony rates of intake of pollen and nectar were not independent. Selection resulted in an increase in the number of pollen collectors and a decrease in the number of nectar collectors in high strain colonies, while the reciprocal relationship occurred in the low strain. High and low strain colonies also demonstrated different diurnal foraging patterns as measured by the changing proportions of returning pollen foragers. High strain colonies of generation 3 contained significantly less brood than did low strain colonies, a consequence of a constraint on colony growth resulting from a fixed nest volume and large quantities of stored pollen. These components represent selectable colony-level traits on which natural selection can act and shape the social organization of honey bee coloniesCommunicated by R.F.A. Moritz     

Vocal mediation of foraging competition in the cooperatively breeding green woodhoopoe (<Emphasis Type="Italic">Phoeniculus purpureus</Emphasis>)

Many group-living species produce frequent vocalisations when foraging, but the function of these food-associated calls is often difficult to divine. I investigated the kek call of the cooperatively breeding green woodhoopoe (Phoeniculus purpureus), a species in which individuals have preferred foraging techniques dependent on their bill size. Individuals called at a greater rate (1) in foraging compared to non-foraging situations, and (2) in groups containing potential foraging competitors (i.e. individuals that foraged using the same preferred techniques). I therefore asked whether the kek call is used to recruit conspecific foragers or whether it acts as a vocal signal of foraging niche and mediates foraging competition. Foragers that were vocalising were no more likely to be approached than those that were silent, and individuals gained no foraging advantage from the close proximity of another group member. Thus, keks are unlikely to be used to recruit conspecifics. Instead, they appear to regulate spacing between potential foraging competitors. Although an individual forager was equally likely to be closely approached by all other group members, it increased its calling rate only in response to potential foraging competitors. This increase in calling rate resulted in the approaching individual moving away, thus maintaining some separation between individuals that forage in the same way. Maintenance of such spacing is important because the success rate of an individual decreased when a foraging competitor was close by.Communicated by M. Leonard