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Species-And Sex-Specific Differences in Foraging Behaviour and Foraging

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Species-And Sex-Specific Differences in Foraging Behaviour and Foraging Vol. 391: 267–278, 2009 MARINE ECOLOGY PROGRESS SERIES Published September 28 doi: 10.3354/meps07981 Mar Ecol Prog Ser Contribution to the Theme Section ‘Spatiotemporal dynamics of seabirds in the marine environment’ OPENPEN ACCESSCCESS Species- and sex-specific differences in foraging behaviour and foraging zones in blue-footed and brown boobies in the Gulf of California Henri Weimerskirch1,*, Scott A. Shaffer2, Yann Tremblay2, Daniel P. Costa2, Hélène Gadenne1, Akiko Kato3, 6, Yan Ropert-Coudert3, 6, Katsufumi Sato4, David Aurioles5 1Centre d’Etudes Biologiques de Chizé, Centre National de la Recherche Scientifique, 79360 Villiers en Bois, France 2Ecology & Evolutionary Biology, University of California Santa Cruz, Santa Cruz, California 95060, USA 3National Institute of Polar Research, 1-9-10 Kaga, Itabashi-ku, Tokyo 173-8515, Japan 4International Coastal Research Center, Ocean Research Institute, The University of Tokyo, 2-106-1 Akahama, Otsuchi, Iwate 028-1102, Japan 5Departamento de Pesquerias y Biologia Marina, CICIMAR-IPN, La Paz, B.C.S., Mexico 6Present address: Institut Pluridisciplinaire Hubert Curien, Centre National de la Recherche Scientifique,67037 Strasbourg, France ABSTRACT: 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; fe- males 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 simi- lar 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 differ- ences 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 differ- ences in observed foraging behaviour were partly or totally explained by body size (flight speeds, for- aging 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. KEY WORDS: Accelerometers · GPS tracking · Sula leucogaster · Sula nebouxii · Area-restricted search · Fractal landscape method · Diet · Isotopes Resale or republication not permitted without written consent of the publisher INTRODUCTION patrically, niche differentiation is expected at equilib- rium. Partitioning of food sources can occur in sym- Understanding how closely related species can patric species by differential selection of foraging coexist has been a long-lasting subject of research (e.g. habitat, foraging strategy or prey choice. Body size Pianka 1981, Ricklefs 1990). When species breed sym- differences between species may also favour niche dif- *Email: [email protected] © Inter-Research 2009 · www.int-res.com 268 Mar Ecol Prog Ser 391: 267–278, 2009 ferentiation (Bowers & Smith 1979, Clutton-Brock et not extensive compared to other species of birds. Nev- al. 1987, Andersson 1994, LeBoeuf et al. 2000). Within ertheless, sex-specific differences in foraging behav- species, body size differences between sexes are com- iour have been found in several species with pro- mon, and the extent of the difference in size varies nounced sexual dimorphism (Weimerskirch et al. 1993, extensively according to the taxa considered (Anders- 2006, Kato et al. 1999, González-Solís et al. 2000a,b, son 1994). Three major hypotheses have been pro- Phillips et al. 2004), but also in species with no size posed to explain the evolution of sexual size dimor- dimorphism (Gray & Hamer 2001, Lewis et al. 2002). phism: (1) sexual selection, (2) intersexual food compe- These latter examples suggest that differences in for- tition, and (3) reproductive role division, and empirical aging behaviour may not always be related to the studies have demonstrated that each of the 3 mecha- maintenance of sexual size dimorphism (Lewis et al. nisms operates in natural populations (Hedrick & 2005). RSD is also found in several seabird families Temeles 1989). Sex differences in food and foraging such as boobies, frigatebirds and skuas. In boobies, sex ecology have often been proposed as important factors differences in foraging behaviour have been found leading to the evolution of size dimorphism between in several species (Lewis et al. 2005, Weimerskirch et sexes (Andersson & Norberg 1981, Shine 1989, Mueller al. 2006, Zavalaga et al. 2007), and the degree of 1990), and many empirical studies have highlighted difference appears to match the extent of sexual di- such differences (e.g. Selander 1966, Schoener 1967, morphism for some foraging parameters such as dive Pierotti 1981, Le Boeuf et al. 2000, Cook et al. 2007). depths or foraging duration (Lewis et al. 2005). Apart from diet, it is often difficult to study foraging In the present study, we examined sex differences in behaviour in many fast-moving or wide-ranging ani- the foraging behaviour of 2 sympatric booby species of mals under natural conditions. Therefore, we generally different body size, each species presenting RSD with lack information on the way sexes or species can differ extensive size dimorphism. We studied brown boobies in their foraging behaviour, such as movement, tech- Sula leucogaster brewsteri and blue-footed boobies niques or effort, which limits our ability to relate forag- S. nebouxii breeding on an island in the Gulf of Cali- ing and body size differences. fornia, using (1) highly accurate miniaturised GPS data In birds, where flight is strongly constrained by loggers to examine the spatial distribution and forag- physics, and in particular by structural size and mass, ing movements of each species and sex, and (2) data body size differences between sexes are less exagger- loggers that measured diving depth and acceleration ated than in mammals and reptiles (Andersson 1994). to study the details of the diving behaviour and time- Although not as prominent, sexual dimorphism occurs budget activity. In addition, diet differences between in many bird taxa where males are larger than study groups were examined by collecting regurgi- females, and the differences have most often been tated stomach contents, and stable isotopes were stud- related to sexual selection. Reversed size dimorphism ied from blood and feather samples. Our primary (referred to as reversed sexual dimorphism, RSD) also objective was to examine whether both species dif- exists among several avian taxa. In species exhibiting fered in their foraging behaviour, and whether forag- sexual size dimorphism, significant differences in for- ing behaviour differed between sexes within each spe- aging behaviour have been found (e.g. Newton 1979). cies. Because one species is smaller than the other, and Within a particular taxonomic family, the extent of sex- in contrast to the approach of Lewis et al. (2005), who ual dimorphism can vary according to species, sug- tested sexual differences within 2 species, one with a gesting that mechanisms leading to the evolution of higher degree of sexual dimorphism than the other, we dimorphic patterns within a family should vary in their chose to investigate differences along a gradient of extent. In the hypothesis that sexual dimorphism is individuals ranging from small male brown boobies, related to foraging behaviour, the extent of niche par- medium-sized female brown and male blue-footed titioning and congruent differences in foraging behav- boobies, and larger female blue-footed boobies. This iour between 2 species are expected to differ in pro- setting allowed us to disentangle the respective roles portion to sexual dimorphism. Thus, when examining of each species–sex combination in foraging behaviour the evolution of size dimorphism and its relationship at this breeding colony, while taking into account the with foraging behaviour, it is of particular interest to influence of size and mass on foraging parameters. compare the respective influence of species, sex and the degree of size difference on the foraging behaviour of closely related species. However, this has rarely MATERIALS AND METHODS been done (e.g. Paredes et al. 2008). In seabirds, males and females have similar roles The study was carried out on Isla San Ildefonso when breeding, plumage characteristics are generally (111.4° W, 26.6° N) in the Gulf of California, Mexico, similar between the sexes, and sexual dimorphism is between 3 and 12 March 2006. San Ildefonso is a 1 km Weimerskirch et al.: Foraging behaviour of boobies 269 long island located ca. 10 km from the
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