Avian daily foraging patterns: Effects of digestive constraints and variability

Summary Birds show a typical daily pattern of heavy morning and secondary afternoon feeding. We investigate the pattern of foraging by a bird that results in the lowest long-term rate of mortality. We assume the following: mortality is the sum of starvation and predation. The bird is characterized by two state variables, its energy reserves and the amount of food in its stomach. Starvation occurs during the day if the bird's reserves fall to zero. The bird starves during the night if the total energy stored in reserves and the stomach is less than a critical amount. The probability that the bird is killed by a predator is higher if the bird is foraging than if it is resting. Furthermore, the predation risk while foraging increases with the bird's mass. From these assumptions, we use dynamic programming techniques to find the daily foraging routine that minimizes mortality. The principal results are (1) Variability in food finding leads to routines with feeding concentrated early in the day, (2) digestive constraints cause feeding to be spread more evenly through the day, (3) even under fairly severe digestive constraints, the stomach is generally not full and (4) optimal fat reserve levels are higher in more variable environments and under digestive constraints. This model suggests that the characteristic daily feeding pattern of small birds is not due to digestive constraints but is greatly influenced by environmental variability.     

When are two heads better than one? Visual perception and information transfer affect vigilance coordination in foraging groups

Animals frequently raise their heads to check for danger. Ina group, individuals generally raise their heads independently.Earlier models suggest that all group members could gain bycoordinating their vigilance, i.e., each member raising itshead when others are not. We re-examine these suggestions, consideringgroups of different sizes, in light of empirical findings that:(1) animals can sometimes detect a predator without raisingtheir heads, and (2) when one member of a group detects a predator,the information does not always spread to other members of thegroup. Including these effects in models generally decreasesthe value of coordinated vigilance. Coordinated vigilance ishighly favored only when animals have a low probability of detectingpredators without lifting their heads but a high probabilityof being warned when another member of the group detects a predator.For other combinations, coordinated vigilance has little valueand may have a negative value. Group size has contrasting effectsdepending on how social information is obtained. Coordinationis favored in smaller groups when one or more detecting individualsprovide a constant amount of information to individuals unawareof the predator. On the other hand, coordination is favoredin larger groups if each detecting individual provides unawareindividuals with an independent source of information (i.e.,if the amount of information increases as the number of detectingindividuals increases). These results depend on the balanceof an escape due to social information and dilution of riskin groups with imperfect information spread. This frameworkcould be tested by examining species with different visual fieldsand in different environments.     

Effects of feral cats on the evolution of anti-predator behaviours in island reptiles: insights from an ancient introduction

Exotic predators have driven the extinction of many island species. We examined impacts of feral cats on the abundance and anti-predator behaviours of Aegean wall lizards in the Cyclades (Greece), where cats were introduced thousands of years ago. We compared populations with high and low cat density on Naxos Island and populations on surrounding islets with no cats. Cats reduced wall lizard populations by half. Lizards facing greater risk from cats stayed closer to refuges, were more likely to shed their tails in a standardized assay, and fled at greater distances when approached by either a person in the field or a mounted cat decoy in the laboratory. All populations showed phenotypic plasticity in flight initiation distance, suggesting that this feature is ancient and could have helped wall lizards survive the initial introduction of cats to the region. Lizards from islets sought shelter less frequently and often initially approached the cat decoy. These differences reflect changes since islet isolation and could render islet lizards strongly susceptible to cat predation.     

Why is Female Choice not Unanimous? Insights from Costly Mate Sampling in Marine Iguanas

Females do not unanimously choose the single 'best' male, even when female choice is strong, such as in leks, or in polygynous mating situations. A possible explanation is that females base their choices on limited information, perhaps because gathering information is costly. We tested this hypothesis by continuously observing individual female marine iguanas throughout the mating period in order to document the information they gathered about each potential mate. Females actively visited approximately five additional males during the 3 d prior to copulation, compared to the males seen on their normal foraging routes. Females were more likely to visit large-bodied males, but preferentially copulated with the male that had the highest display rate of all males they visited. Females that mated on a dense territory cluster mated with more active males than did those that mated on dispersed territories. However, females on a dense cluster also lost more body mass, potentially as a consequence of high rates of interaction with males. This mass loss may represent an important cost and result from postural changes in response to male attention. Such costs may explain why females only gather a certain amount of information and why females on dispersed territories choose less active mates. Lack of complete information introduces subjectivity into female choice: what is perceived as best by one female may not be perceived as best by another. Thus, lack of complete information may prevent unanimity of female choice.     

Mutualism among safe, selfish sentinels: a dynamic game

Sentinels are group members that watch from prominent positions. Sentinel interchanges often appear orderly, and groups with sentinels rarely have zero or many sentinels. A dynamic game was constructed to examine if these observations about sentinels could be based on selfish actions by individual group members. In this game, each group member chose to forage or be a sentinel based on its own energetic state and the actions of others. Sentinels received a selfish antipredator benefit if their ability to detect approaching predators more than compensated for their increased exposure to undetected predators. Provided sentinels were relatively safe and that detection information spread to other group members when sentinels detected predators, sentinels appeared highly coordinated for all combinations of parameters. This apparent coordination was based on mutualism because each individual gained by being a sentinel when other group members were not (and foraging when other group members were being sentinels). The model was very robust, but exact level of sentinel behavior varied somewhat with changes in foraging and predation parameters. This model could best be tested by testing its assumptions about sentinel safety, foraging-predation trade-offs, and information transfer in groups.     

Florida Scrub-Jays Compensate for the Sentinel Behavior of Flockmates

Sentinel coordination requires that individuals react to the sentinel behavior of others. Previous work showed that Florida scrub‐jays are sentinels more often when given supplemental food. Here we measured how birds in pairs reacted when their mates were fed. Scrub‐jays were sentinels less when their mates were fed, demonstrating compensation. Indirect evidence suggests that this compensatory decrease in sentinel behavior was smaller than the increase in sentinel behavior by their mates. In addition, males in newly established groups were sentinels less often.     

Risk allocation and competition in foraging groups: reversed effects of competition if group size varies under risk of predation

Animals often feed more quickly when in larger groups. This group-size effect is often explained by safety advantages for groups but an alternative explanation is that animals feed faster in larger groups because of greater scramble competition for limited food. We show that predation risk enhances the group-size effect if groups vary in size. By contrast, competition leads to the group-size effect only when individuals feed in groups of constant size. When individuals feed in groups that vary in size, the best strategy for dealing with competition is to feed intensely when in smaller groups and feed little when in larger (more competitive) groups. In all situations, the effects of competition interact with the effects of predation risk in a simple multiplicative way. Our results suggest that scramble competition is not a general explanation for the group-size effect on vigilance in situations where group size changes relatively rapidly.