When 2 closely related species co-occur, each exhibiting sex-specific differences in size, resource partitioning is expected. We studied sex-specific foraging behaviour of 2 sympatric seabird species in the Gulf of California to disentangle the respective influence of species and sex, but also mass and size of individuals, on observed foraging behaviour. We used highly accurate data loggers to study movements, diving behaviour and activity of brown and blue-footed boobies rearing young chicks. Interspecific differences were limited; brown boobies had longer foraging trips and spent less time on the water than blue-footed boobies. The major differences observed were sex-specific; females of each species tended to have longer foraging trips, foraged farther from the colony, flew greater distances and had larger zones of area-restricted search. These sex-specific differences were more prominent in brown than in blue-footed boobies. Diet and stable isotope analyses showed that, during the study period, both species fed mainly on sardines, at similar trophic levels and in similar zones; outside the breeding season, the carbon and nitrogen signatures from feathers were also similar on average. In these sympatric species that feed on a superabundant prey, sex-specific differences appear to have a greater role than species-specific differences. We suggest that sex-specific differences may be mainly related to breeding involvement, as males are more involved in nest attendance and defence and females are greater provisioners. However, we show that several sex-specific differences in observed foraging behaviour were partly or totally explained by body size (flight speeds, foraging range, flapping frequency) or by body mass (depths attained during diving, duration of dives), which are parameters influenced by biomechanical constraints such as flight and diving.
Reversed sexual dimorphism (RSD) may be related to different roles in breeding investment and/or foraging, but little information is available on foraging ecology. We studied the foraging behaviour and parental investment by male and female masked boobies, a species with RSD, by combining studies of foraging ecology using miniaturised activity and GPS data loggers of nest attendance, with an experimental study where flight costs were increased. Males attended the chick more often than females, but females provided more food to the chick than males. Males and females foraged during similar periods of the day, had similar prey types and sizes, diving depths, durations of foraging trips, foraging zones and ranges. Females spent a smaller proportion of the foraging trip sitting on the water and had higher diving rate than males, suggesting higher foraging effort by females. In females, trip duration correlated with mass at departure, suggesting a flexible investment through control by body mass. The experimental study showed that handicapped females and female partners of handicapped males lost mass compared to control birds, whereas there was no difference for males. These results indicate that the larger female is the main provisioner of the chick in the pair, and regulates breeding effort in relation to its own body mass, whereas males have a fixed investment. The different breeding investment between the sexes is associated with contrasting foraging strategies, but no clear niche differentiation was observed. The larger size of the females may be advantageous for provisioning the chick with large quantities of energy and for flexible breeding effort, while the smaller male invests in territory defence and nest guarding, a crucial task when breeding at high densities. In masked boobies, division of labour appears to be maximal during chick rearin-g-the most energy-demanding period--and may be related to evolution of RSD.
Understanding factors shaping the spatial distribution of animals is crucial for the conservation and management of wildlife species. However, few studies have investigated density-dependent habitat selection in wild populations in non-equilibrium conditions and at varying spatial scales. Here, we investigated densitydependent habitat selection at varying spatial scales in an increasing white stork Ciconia ciconia population using a long term data set in western France. During a 16-year study period, the breeding population density increased from 0.66 nests per 100 km 2 to 6.6 nests per 100 km 2 .A t the beginning of the colonisation of the area settlement probability of storks was mainly positively affected by grasslands located near wetlands. Areas with intensive or moderately intensive agriculture were extremely unlikely to be occupied by breeding birds. However, selection for the initially preferred habitats faded as stork density increased although the proportions of habitat types remained unchanged. At the same time selection for initially less favoured habitats had increased. Moreover, we found that the spatial scale of selection for each foraging habitat variable was consistent over time. Our results suggest that snapshot analyses of resource selection in populations at high density may be misleading for population conservation or management. In contrast, a longitudinal approach to resource selection can be a valuable tool for understanding resource limitation.
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