Begging and provisioning in broods of asynchronously-hatched yellow-headed blackbird nestlings

Studies of begging have found a positive relationship between begging level and provisioning level. Studies of unequal nestlings, however, have found that small nestlings generally beg more but are fed less than their larger siblings. We manipulated the begging levels of yellow-headed blackbird (Xanthocephalus xanthocephalus) chicks to investigate how begging benefits individuals in broods of unequal siblings. Food-deprived chicks begged more and were fed more; satiated chicks begged less and were fed less. When we deprived each chick of a brood in turn, large and small chicks generally increased begging and received more provisioning. Small chicks, however, rarely received more food than their larger siblings even when they behged relatively more. Parent yellow-headed blackbirds increase provisioning to hungry begging chicks, but also allocate food based on relative offspring size.     

Net energy versus efficiency maximizing by foraging ring-billed gulls

Summary Ring-billed gulls (Larus delawarensis) breeding at Dog Lake, Manitoba often feed by following tractors pulling cultivating implements around fields. Tractor-following gulls always land immediately behind the cultivating implement, where they feed on earthworms or grain. Afeer a feeding bout on the ground (patch residence time), gulls fly up, pursue the tractor and repeat the cycle. We use net energy maximizing (energy gained per unit time) and efficiency maximizing (energy gained per unit energy expended) models to make quantitative predictions of patch residence time, and compare these predictions to observations. If we assume flight speed to be constrained at the observed mean, the observations fall between the predictions of the two models, and the models explain approximately equal and highly significant proportions of the overall variation in patch residence time. If both flight speed and patch residence time are allowed to vary in the models, the efficiency maximizing model more closely predicts observed patch residence times, and the net energy maximizing model more closely predicts observed flight speeds. We discuss whether breeding ringbilled gulls may be truly intermediate between net energy and efficiency maximizing, and how measurements of flight speed may be useful in further investigations.     

Body condition and the grouping behavior of brood-caring female common eiders (<Emphasis Type="Italic">Somateria mollissima</Emphasis>)

Both theoretical and empirical work has shown that group size increases with increasing ecological constraints on solitary breeding. Ecological constraints refer to extrinsic factors such as availability of breeding sites, food or mates. Common eider (Somateria mollissima) females pool their broods and share brood-rearing duties, or rear broods alone. Females are often in poor condition at hatching, as incubation is accomplished without feeding, and variation in body condition is largely environmentally induced and thus unpredictable. We found that the intensity of and duration of parental care that females provide is positively correlated with their body condition at hatching. This suggests that body condition is an ecological constraint on successful solitary breeding. We further observed that group productivity in common eider broods is a decelerating function of the number of tending females. As predicted, females in poorer condition (i.e., facing stronger ecological constraints) were found in larger groups. This result is straightforward if solitary tenders can enter any group at no cost. However, if entry is group-controlled, stable groups of non-relatives are predicted not to occur when per capita reproduction declines with group size. The N-person staying incentive model permits groups to form under these conditions, because reproduction is unevenly divided between dominants and subordinates in the group. We discuss the plausibility of these alternative models of group size for understanding the grouping behavior of brood-caring female common eiders.Communicated by M. Webster     

Condition-dependent response to changes in pollen stores by honey bee (Apis mellifera) colonies with different parasitic loads

The impact of a parasitic infestation may be influenced by nutritional state, in both individuals and colonies. This study examined the interaction between pollen storage and the effects of an infestation by the mite, Varroa jacobsoni Oudemans, in colonies of the honey bee, Apis mellifera L. We manipulated the pollen storage and mite infestation levels of colonies, and measured pollen foraging and brood rearing. Increased pollen stores decreased both the number of pollen foragers and pollen load size, while initially at least foragers from colonies with moderate infestations carried smaller pollen loads than those from lightly infested colonies. Over the course of the experiment, all colonies significantly increased pollen-foraging rates and pollen consumption, which was presumably a seasonal effect. Lightly infested colonies exhibited a larger increase in pollen forager number than moderately infested colonies, suggesting that more intense mite infestations compromised forager recruitment. Brood production was not affected by the addition of pollen, but moderately infested colonies were rearing significantly less brood by the end of the experiment than lightly infested colonies. Furthermore, the efficiency with which colonies converted pollen to brood decreased as the pollen storage level decreased and the infestation level increased. The results of this study may indicate that honey bee colonies adaptively alter brood-production efficiency in response to parasitic infestations and seasonal changes. Received: 3 May 1999 / Received in revised form: 14 September 1999 / Accepted: 25 September 1999     

The conflict between feeding and territorial defence in the great tit

Summary A field experiment investigated whether feeding and territorial defence competed for time in the activity budget of territorial male great tits during the spring. Feeding tables were placed on the territories of five males. Five other territorial males also fed at these tables, while ten additional males had no access to any feeding tables. A standardised intrusion using playback of territorial song and a stuffed mount of a male great tit was conducted on each of the twenty territories, and the response of the resident male measured. All of the males with access to a feeding table responded more vigorously to the intrusion than the males who received no extra food. These results are not attributable to the fact that males defended their territories more vigorously because of the addition of feeding tables, since males that gained extra food outside the boundaries of their own territories also defended more vigorously than birds with no access to a feeding table. The results support the conclusion that the provisioned males defended more vigorously because they could afford to take more time out from feeding.     

The effects of wing loading and gender on the escape flights of least sandpipers (<Emphasis Type="Italic">Calidris minutilla</Emphasis>) and western sandpipers (<Emphasis Type="Italic">Calidris mauri</Emphasis>)

High body mass caused by fat storage during migration is believed to increase a bird's risk of predation by decreasing its ability to escape predators. We demonstrate the negative effect of wing loading (mass/wing area) on escape speed and angle of two migrating species of shorebird. We also show significant differences in escape performance between the species and genders. To help explain these differences, we test two potential proximate causes, wing shape and leg bone length. Wing shape is correlated with differences in escape performance between the species, but we found no correlation of wing shape or leg bone length with gender. Ultimately, greater predation risk due to habitat use or larger body size, for the species and genders respectively, may have resulted in evolution of enhanced escape ability.     

Intensity of territorial defense in red squirrels: an experimental test of the asymmetric war of attrition

Summary Red squirrels defend exclusive, individual territories year round, 20% to 50% of females do not breed in any given year, and breeding females raise juveniles on their territories. Breeding is asynchronous, and the offspring of early-breeding females are more likely to hold an independently won territory than are late-born offspring. Based on the asymmetric war of attrition, we made the following predictions: (1) squirrels would respond more intensely to the calls of unfamiliar individuals than to the calls of neighbors; (2) breeding females would respond more intensely to unknown calls than would non-breeding females or males; (3) early-breeding females would respond more intensely than would late-breeding females to unknown calls; and (4) all classes of squirrels would respond similarly to the calls of neighbors. Playback experiments supported the predictions. Alternative hypotheses of kin selection, risk of infanticide, and seasonal difference in intruder pressure could not explain the results.     

Interannual differences in the relative timing of southward migration of male and female western sandpipers (Calidris mauri)

In the shorebird subfamily Calidridinae, one of the parents shortens parental care and initiates southward migration before the other. We estimated the difference in passage date between male and female western sandpipers (Calidris mauri) at their first major stopover on the southward migration from breeding areas in Alaska, in 18 years between 1978 and 2000. Overall, adult females preceded adult males by 1.22 days. A novel finding was that among juveniles, which migrate approximately a month later than adults, females preceded males by similar magnitude (1.14 days). There was wide variation among years, however, and males actually preceded females in years with late hatch. We relate these findings to hypotheses for female-first southward migration in sandpipers.     

Differential predator escape performance contributes to a latitudinal sex ratio cline in a migratory shorebird

Sexual segregation outside the mating season is common in vertebrates, and has been attributed to sexual differences in predator escape performance in ungulates and fish, but not in birds. Here, we tested the hypothesis that sex- and latitude-specific predator escape performance underlies the differential nonbreeding distribution of western sandpipers (Calidris mauri), a migratory shorebird. Females overwinter further south along the American Pacific coast, creating a latitudinal cline in sex ratio. Escape performance is reduced with increasing body mass, and birds generally carry less fat further south. Western sandpipers with poor escape performance were therefore predicted to prefer southern sites to reduce the risk of mortality posed by predators. Data from four nonbreeding latitudes showed that wing loading, used as an index of escape performance, was overall higher for females, and that it decreased with latitude in both sexes. Within latitudes, wing loading was lower at smaller, and presumably more dangerous, sites. Flight response to a predatory attack was longer in the south. Mortality risk offers a novel and candidate explanation for differential distribution patterns in western sandpipers and possibly other avian migrants.     

Mortality-minimizing sandpipers vary stopover behavior dependent on age and geographic proximity to migrating predators

Ecological theory for long-distance avian migration considers time-, energy-, and mortality-minimizing tactics, but predictions about the latter have proven elusive. Migrants must make behavioral decisions that can favor either migratory speed or safety from predators, but often not both. We compare the behavior of adult and juvenile western sandpipers Calidris mauri during the course of their temporally segregated passages at a major stopover site. Here, the passage and winter arrival of an important predator, the peregrine falcon Falco peregrinus begins near the end of the adult sandpiper passage (July) and increases rapidly through the juvenile passage (August). The mortality-minimizing hypothesis predicts that as the falcon front is distant but approaching, sandpipers should initially increase the fuel-loading rate (lowered vigilance and predator apprehension) to increase migration speed and so maintain their head start. As the falcon front gains proximity to and passes over the stopover site, sandpipers should become increasingly cautious. Our measurements show that adults decreased vigilance during the period prior to falcon arrival, and had lower vigilance overall than juveniles. Juveniles were more apprehensive, flying further and longer in response to disturbance by a falcon silhouette. This trend was reversed in response to a human approach. Both groups were more vigilant and more apprehensive in a study year with earlier falcon arrival. These results suggest that late (juvenile) and early (adult) migrants minimize mortality on migration in different ways, adults by increased migratory speed at the expense of caution on stopover sites, and juveniles by increased caution at the expense of speed.