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Monday, 17 March 2014

Journal of Avian Biology - 'Seabird' Virtual Issue March 2014: Abstract Listing

Introduction from the Editors
"With this Virtual Issue of Journal of Avian Biology we wish to highlight some of the many papers focusing on Seabird biology published by the Journal. The issue is especially compiled for the International Seabird Group Conference in Oxford in March 2014 and represent the diverse and complex biology of this fascinating group of birds. Journal of Avian Biology has a strong tradition of publishing papers on all aspects of seabird biology and we hope that you will find this thematic virtual issue especially interesting. All the papers are freely available for everyone to enjoy."

Below are the links, titles, authors, and abstracts of the free access papers.

The individual counts: within sex differences in foraging strategies are as important as sex-specific differences in masked boobies Sula dactylatra
Julia Sommerfeld, Akiko Kato, Yan Ropert-Coudert, Stefan Garthe, Mark A. Hindell

Understanding how animals allocate their foraging time is a central question in behavioural ecology. Intrinsic factors, such as body mass and size differences between sexes or species, influence animals’ foraging behaviour, but studies investigating the effects of individual differences in body mass and size within the same sex are scarce. We investigated this in chick-rearing masked boobies Sula dactylatra, a species with reversed sexual dimorphism, through the simultaneous deployment of GPS and depth-acceleration loggers to obtain information on foraging movements and activity patterns. Heavier females performed shorter trips closer to the colony than lighter females. During these shorter trips, heavier females spent higher proportions of their flight time flapping and less time resting on the water than lighter females did during longer trips. In contrast, body mass did not affect trip duration of males, however heavier males spent less time flapping and more time resting on the water than lighter males. This may occur as a result of higher flight costs associated with body mass and allow conservation of energy during locomotion. Body size (i.e. wing length) had no effect on any of the foraging parameters. Dive depths and dive rates (dives h−1) were not affected by body mass, but females dived significantly deeper than males, suggesting that other factors are important. Other studies demonstrated that females are the parent in charge of provisioning the chick, and maintain a flexible investment under regulation of their own body mass. Variation in trip length therefore seems to be triggered by body condition in females, but not in males. Consequently, shorter trips are presumably used to provision the chick, while longer trips are for self-maintenance. Our findings underline the importance of accounting for the effects of body mass differences within the same sex, if sex-specific foraging parameters in dimorphic species are being investigated.
Networks of prey specialization in an Arctic monomorphic seabird
Jennifer F. Provencher, Kyle H. Elliott, Anthony J. Gaston, Birgit M. Braune

Generalist predator populations are sometimes made up of individuals that specialize on particular prey items. To examine specialization in thick-billed murres Uria lomvia during self-feeding we obtained stomach contents and muscle stable isotope values for 213 birds feeding close to five colonies in the Canadian Arctic. Adults were less specialized during self-feeding than during chick-provisioning. Nonetheless, particular specialists clustered together within the foraging network. While sexes showed similar levels of specialization, individuals of the same sex clustered together within the foraging network. The significant degree of clustering regardless of sex showed that individuals specializing on one prey item tend to also specialize on another, although network topology varied from colony to colony. Adult muscle stable isotope values correlated with the stable isotope values of the prey found in stomachs, at least at the one colony with relevant prey data, suggesting that specializations are maintained over time. Degree of specialization increased with niche width across the five colonies, but similarity in gastro-intestinal and bill morphology was independent of dietary similarity. Thus, although individual specialization is thought to play a key role in sympatric speciation through trophic specialization, we found no support for an association between morphology and foraging patterns in our species. We conclude that self-feeding murres show clustered dietary specialization, and that specialization is highest where diet is most diverse.
Movements and activities of male black-tailed gulls in breeding and sabbatical years
Kentaro Kazama, Kazuhiko Hirata, Takashi Yamamoto, Hiroshi Hashimoto, Akinori Takahashi, Yasuaki Niizuma, Philip N. Trathan, Yutaka Watanuki

Long-lived animals sometimes skip one or more breeding seasons; however, little is known about their movements and activities during such ‘sabbatical’ periods. Here we present novel data on year-round movements and activities of two male black-tailed gulls Larus crassirostris during a sabbatical year. We compare the data with those in a year when they bred and with those of two other breeding males. The year-round migration routes of two sabbatical males were consistent with those of the breeding males: they returned to the breeding area but did not visit the colony in the sabbatical year. They landed more frequently on water (a potential index of foraging effort) during the non-breeding autumn and winter prior to the sabbatical year than before breeding. Sabbatical gulls may forage more intensively to recover body condition immediately after breeding.
Colouration in Atlantic puffins and blacklegged kittiwakes: monochromatism and links to body condition in both sexes
Claire Doutrelant, Arnaud Grégoire, Doris Gomez, Vincent Staszewski, Emilie Arnoux, Torkild Tveraa, Bruno Faivre, Thierry Boulinier

Sexual dimorphism is widely used as an indirect measure of the intensity of sexual selection. It is also a way to evaluate whether different selective pressures act on males and females. Dichromatism, defined as a difference in colouration between males and females, may for instance result from selection for crypsis in females and selection for conspicuousness in males. Here, we conducted a study to investigate whether differential sexual selective pressures might act on the colour traits of two colonial seabird species, the Atlantic puffin Fratercula artica and the black-legged kittiwake Rissa tricactyla. First, we used spectrophotometry and visual modelling to determine whether these presumed monomorphic birds are really monochromatic from an avian perspective (birds and humans have a different vision). Second, we estimated whether some of their colourations have the potential to be sexually or socially selected by determining whether these colourations were related to body condition in males and females, and whether the yellow, orange and red colourations may contain carotenoid pigments. Our results indicated that both species were fully monochromatic from an avian perspective. Moreover, our preliminary analyses suggested that the yellow, orange and red colours of these birds contained carotenoids. Lastly, some indices of colouration were positively linked to estimates of condition. Birds in better condition had redder gape (both species) and bill (puffins). In puffins, the relation between condition and gape colouration was significantly stronger in females than males. By contrast, the size of the gape rosette was larger in males than females. The positive links we found between colour indices and condition, together with the absence of sexual dichromatism, suggest that mutual sexual selection may act in these two species.
Chronic stress in infancy fails to affect body size and immune response of adult female blue-footed boobies or their offspring
María Cristina Carmona-Isunza, Alejandra Núñez-de la Mora, Hugh Drummond

Experiments on birds, fish and mammals have shown that adverse conditions during infancy can produce diverse long-term and delayed deficits during adulthood, prejudicing both the individual and its offspring. Natural selection should prepare animals to cope with adversity of the type, magnitude and timing that commonly occur in their natural habitat, but very little is known about such evolved developmental buffering against natural ‘poor starts’ in life. In two-chick broods of the blue-footed booby Sula nebouxii, the junior (younger) chick usually experiences aggressive subordination, reduced nutrition and growth and elevated circulating corticosterone. To test whether this poor start produces long-term, delayed or intergenerational deficits in body size, body condition or cell-mediated immune response, we measured 3–8 yr old female breeders banded as chicks, and their infant offspring. Results failed to support our predictions. Compared to former seniors and former singletons (solitary nestlings), former juniors showed no deficit in cell-mediated immune response at any age. They showed an 8.04% deficit in body condition at age 4–6 yr but this deficit disappeared completely by age 7–8 yr. Furthermore, their offspring showed no deficits in body size, body condition or immune response. Junior chicks are affected by their poor start, but their developmental resilience, also confirmed by studies of post-fledging survivorship, recruitment, natal dispersal, aggressive nest defense and reproduction, is evidence of evolved developmental buffering against predictable adversity during infancy.
Energetic consequences of contrasting winter migratory strategies in a sympatric Arctic seabird duet
Jérôme Fort, Harald Steen, Hallvard Strøm, Yann Tremblay, Eirik Grønningsæter, Emeline Pettex, Warren P. Porter, David Grémillet

At the onset of winter, warm-blooded animals inhabiting seasonal environments may remain resident and face poorer climatic conditions, or migrate towards more favourable habitats. While the origins and evolution of migratory choices have been extensively studied, their consequences on avian energy balance and winter survival are poorly understood, especially in species difficult to observe such as seabirds. Using miniaturized geolocators, time-depth recorders and a mechanistic model, we investigated the migratory strategies, the activity levels and the energy expenditure of the closely-related, sympatrically breeding Brünnich's guillemots Uria lomvia and common guillemots Uria aalge from Bjørnøya, Svalbard. The two guillemot species from this region present contrasting migratory strategies and wintering quarters: Brünnich's guillemots migrate across the North Atlantic to overwinter off southeast Greenland and Faroe Islands, while common guillemots remain resident in the Barents, the Norwegian and the White Seas. Results show that both species display a marked behavioural plasticity to respond to environmental constraint, notably modulating their foraging effort and diving behaviour. Nevertheless, we provide evidence that the migratory strategy adopted by guillemots can have important consequences for their energy balance. Overall energy expenditure estimated for the non-breeding season is relatively similar between both species, suggesting that both southward migration and high-arctic winter residency are energetically equivalent and suitable strategies. However, we also demonstrate that the migratory strategy adopted by Brünnich's guillemots allows them to have reduced daily energy expenditures during the challenging winter period. We therefore speculate that ‘resident’ common guillemots are more vulnerable than ‘migrating’ Brünnich's guillemots to harsh winter environmental conditions.
Specialization to cold-water upwellings may facilitate gene flow in seabirds: new evidence from the Peruvian pelican Pelecanus thagus (Pelecaniformes: Pelecanidae)
Will S. Jeyasingham, Scott A. Taylor, Carlos B. Zavalaga, Alejandro Simeone, Vicki L. Friesen

Recent research has shown that tropical seabirds specialized to feed on cold water upwellings exhibit low population genetic differentiation and high gene flow across large geographic distances. This pattern is opposite to the general pattern of differentiation reported for tropical seabirds, and led us to hypothesize that specialization to cold-water upwellings facilitates gene flow between colonies. As a test of this hypothesis we characterized population differentiation and gene flow across the range of the Peruvian pelican Pelecanus thagus, an upwelling specialist endemic to the Humboldt Current, using an 838 base pair segment of the mitochondrial control region and seven microsatellite loci. In support of our hypothesis we report genetic panmixia across the geographic range of this species and inferred high gene flow between colonies. The high dispersal propensity of upwelling specialist seabirds (adults and/or juveniles) may reduce loss of genetic diversity during population declines, and increase the ability of these species to colonize new islands.
The population increase of common guillemots Uria aalge on Skomer Island is explained by intrinsic demographic properties
Jessica Meade, Ben J. Hatchwell, Julia L. Blanchard, Tim R. Birkhead

The population of common guillemots Uria aalge on Skomer Island, Wales has been monitored since 1963, and in the last 30 yr has increased at an almost constant rate of 5% yr−1. A previous attempt to model the population based on intrinsic demographic parameters estimated over just five years failed to explain the observed population increase, probably because the estimate of juvenile survival was too low. This raised the possibility that immigration fuelled the population increase. Here we use > 30 yr of detailed field observations to re-estimate key population parameters (productivity, adult survival and juvenile survival) in order to model the population. We show that the observed rate of increase can be explained by these intrinsic parameters, and we therefore conclude that immigration is not necessary to generate the observed population growth.
Rates and consequences of relaying in little auks Alle alle breeding in the High Arctic an experimental study with egg removal
Dariusz Jakubas, Katarzyna Wojczulanis-Jakubas

Most seabirds have a small clutch size. Thus, replacement of a clutch after loss can make important contributions to an individual’s lifetime reproductive success. However, in the condition of short polar summer, relaying propensity may be time-constrained. In this study, we investigated rates and consequences of relaying in a small High Arctic seabird, the little auk Alle alle. We performed an experiment in which we removed the single egg from 20 nests of early-laying breeders. We measured relaying rates, and compared chick body mass and breeding success between the experimental and control nests. Despite the narrow window of the Arctic summer and the closely synchronized breeding, 75% of females produced a replacement egg just 2.7% smaller in volume than the first egg. This indicates that in little auks, the demographic effects of disruptions to breeding attempts (by predators, adverse weather or human activity) may be mitigated to some extent by replacement clutches. However, peak body mass and fledging body mass were lower in the experimental than the control chicks. This effect was rather a consequence of late hatching – chicks from replacement clutches followed seasonal decline in peak body mass and fledging mass. Finally, breeding success and chick survival up to 20 d in the experimental nests were respectively 34 and 37% lower than in the control nests. Thus, the quality and post-fledging survival of chicks from the replacement clutches were probably lower compared to the chicks hatched from the first-laid eggs.
Predator–prey interactions between the South Polar skua Catharacta maccormicki and Antarctic tern Sterna vittata
Karel Weidinger, Václav Pavel

Antarctic terns have to co-exist in a limited space with their major nest predator, the skuas. We conducted artificial nest experiments to evaluate the roles of parental activity, nest location and nest and egg crypsis in this simple predator–prey system. Predation on artificial (inactive) nests was higher in traditional nesting sites than in sites previously not occupied by terns, which suggests that skuas memorized past tern breeding sites. Predation on artificial nests in inactive colonies was higher than in active (defended) colonies. Parental defense reduced predation in colonies to the level observed in artificial nests placed away from colonies. This suggests that communal defense can balance the costs of attracting predators to active colonies. Within colonies, predation was marginally higher on experimental eggs put in real nests than on bare ground. Although it seems that the presence of a nest is costly in terms of increased predation, reductions in nest size might be constrained by the need for protective nest structures and/or balanced by opposing selection on nest size. Predation did not differ markedly between artificial (quail) and real tern eggs. A simultaneous prey choice experiment showed that the observed predation rates reflected egg/nest detectability, rather than discrimination of egg types. In summary, nesting terns probably cannot avoid being detected, and they cannot defend their nest by attending them. Yet, by temporarily leaving the nest, they can defend it through communal predator mobbing, and at the same time, they can benefit from crypsis of unattended nest and eggs.
Evidence for strong assortative mating, limited gene flow, and strong differentiation across the blue-footed/Peruvian booby hybrid zone in northern Peru
Scott A. Taylor, David J. Anderson, Carlos B. Zavalaga and, Vicki L. Friesen

Hybrid zones represent natural laboratories in which the processes of divergence and genetic isolation can be examined. The generation and maintenance of a hybrid zone requires mispairing and successful reproduction between organisms that differ in one or more heritable traits. Understanding the dynamics of hybridization between two species requires an understanding of the extent to which they have diverged genetically, the frequency of misparing and hybrid production, and the extent of introgression. Three hundred and twenty one blue-footed Sula nebouxii and Peruvian S. variegata boobies from the eastern tropical Pacific Ocean were analyzed using 19 putatively neutral genetic markers to evaluate interspecific differentiation, to classify morphological hybrids using Bayesian assignments, and to characterize hybridization using cline theory and Bayesian assignments. The species were well differentiated at mitochondrial and nuclear microsatellites, the hybrid zone was bimodal (contained a high frequency of each parental species but a low frequency of hybrids), and morphologically intermediate individuals were most likely F1 hybrids resulting from mating between female Peruvian boobies and male blue-footed boobies. Clines in allele frequency could be constrained to share a common geographic centre but could not be constrained to share a common width. Peruvian and blue-footed boobies hybridize infrequently, potentially due to strong premating reproductive isolation; however, backcrossing appears to facilitate introgression from blue-footed to Peruvian boobies in this hybrid system.
Divergent diving behavior during short and long trips of a bimodal forager, the little auk Alle alle
Zachary W. Brown, Jorg Welcker, Ann M. A. Harding, Wojciech Walkusz, Nina J. Karnovsky

The purpose of this study was to characterize for the first time seabird diving behavior during bimodal foraging. Little auks Alle alle, small zooplanktivorous Alcids of the High Arctic, have recently been shown to make foraging trips of short and long duration. Because short (ST) and long trips (LT) are thought to occur in different locations and serve different purposes (chick- and self-feeding, respectively) we hypothesized that foraging differences would be apparent, both in terms of water temperature and diving characteristics. Using Time Depth Recorders (TDRs), we tested this hypothesis at three colonies along the Greenland Sea with contrasting oceanographic conditions. We found that diving behavior generally differed between ST and LT. However, the magnitude of the disparity in diving characteristics depended on local foraging conditions. At the study site where conditions were favorable, diving behavior differed only to a small degree between LT and ST. Together with a lack of difference in diving depth and ocean temperature, this indicates that these birds did not increase their foraging effort during ST nor did they travel long distances to seek out more profitable prey. In contrast, where local foraging conditions were poor, birds increased their diving effort substantially to collect a chick meal during ST as indicated by longer, more U-shaped dives with slower ascent rates and shorter resting times (post-dive intervals and extended surface pauses). In addition, large differences in diving depth and ocean temperature indicate that birds forage on different prey species and utilize different foraging areas during LT, which may be up to 200 km away from the colony. Continued warming and deteriorating near-colony foraging conditions may have energetic consequences for little auks breeding in the eastern Greenland Sea.
Seasonal variation in pre-fledging survival of lesser scaup Aythya affinis: hatch date effects depend on maternal body mass
Kirsty E. B. Gurney, Robert G. Clark, Stuart M. Slattery

Among temperate-breeding birds, offspring survival and reproductive success are often inversely related to timing of breeding. The mechanisms that produce seasonal declines in offspring survival are not fully understood but may be related to temporal changes in parental quality, environmental quality, or both. We analyzed data for lesser scaup Aythya affinis to evaluate hypothesized effects of parental quality and date on pre-fledging survival. Maternal quality, as indexed by body mass, did not have an independent effect on offspring survival in this species. Maternal body mass did not decline seasonally and did not have an independent effect on duckling survival. Although we did not detect an independent effect of hatch date on duckling survival, duckling survival declined seasonally for broods raised by lightweight females, indicating an interactive effect of maternal mass and date. We hypothesize that this interaction may be driven by seasonally declining food resources coupled with the influence of female condition on the ability to monopolize food resources or remain attentive to the brood. We also tested morphological predictions of the date hypothesis by examining physical characteristics of ducklings. When corrected for age and size, late-hatched ducklings tended to have marginally larger digestive systems and smaller leg muscles than did early-hatched birds. Abundances of intestinal parasites acquired through diet decreased marginally in late-hatched ducklings. Results for digestive system and parasite infection patterns suggested that later-hatched broods may shift diets, consistent with a contribution of environmental factors to seasonal variation in offspring survival. Taken together, our results suggest that both female attributes and environmental conditions may influence seasonal patterns of offspring survival in this species.

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