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Morphological variation in biological structures may be driven by genetic and environmental factors, such as inter- and intraspecific competition for resources. In seabirds, although the bill is also involved in vocalization, olfaction, sexual selection and defence, the main drivers of high morphological plasticity in bill size and shape appear to relate primarily to diet and thus to niche differentiation. Here, we combined geometric morphometrics and comparisons of linear measurements as a precise tool for measuring shape variation in anatomical features, to investigate the differences among species and populations (island groups) in bill shape of three planktivorous petrels (Antarctic prion Pachyptila desolata, blue petrel Halobaena caerulea and thin-billed prion Pachyptila belcheri). Fieldwork was carried out in South Georgia (54° 0' S, 38° 3' W), Falkland (51° 42' S, 57° 51' W), Diego Ramírez (56° 31' S, 68° 44' W) and Kerguelen (49° 20' S, 69° 20' E) Islands, from 2010 to 2021. Results show that the bills of Antarctic prions were more robust and shorter, appropriate for filtering large amounts of small prey. Blue petrels and thin-billed prions had narrower and longer bills, effective for catching and tearing large single prey. Also, Antarctic prions and blue petrels from Kerguelen had longer and narrower bills than conspecifics from other colonies, which could potentially be explained by geographic variation in diet. In conclusion, prey availability and diversity appear to be important factors influencing variation in bill morphology. This study highlights the utility of geometric morphometrics for investigating bill shape variation in seabirds. Nevertheless, further studies are needed to better understand selective pressures leading to morphological variation of biological structures., Author(s): Maria I. Laranjeiro [sup.1] [sup.2] , Marc Farré [sup.1] [sup.3] , Richard A. Phillips [sup.4] , Petra Quillfeldt [sup.5] , Francesco Bonadonna [sup.6] , Charlene Gémard [sup.6] , Maximiliano [...]

Northern Macronectes halli and southern giant petrels M. giganteus are opportunistic predators and the dominant avian scavengers in sub-Antarctic and Antarctic ecosystems. At South Georgia, there are globally important breeding populations of both species; however, no detailed diet study has been carried out at this site for > 35 years. Here, we analysed stomach contents of northern (n = 81) and southern giant petrel (n = 73) chicks at Bird Island, South Georgia (2014/15-2016/17). Specifically, we investigated: (1) interspecific and sexual dietary segregation; (2) diet changes within and among recent years; and (3) long-term diet changes since 1979/80-1980/81. Overall diet composition was similar between species, with penguins comprising approximately half the diet by mass. In both species (but more pronounced in southern giant petrels), prey delivered by female parents included higher proportions (by mass) of penguins and Antarctic krill Euphausia superba, and by male parents included more Antarctic fur seal Arctocephalus gazella carrion and seabirds other than penguins. Consumption of penguins increased, and of seal carrion declined, as chick-rearing progressed, mirroring the decreasing availability of the latter after the peak pupping period in December at South Georgia. Comparisons with data from 1979/80-1980/81 suggest some changes in giant petrel diets; however, these differences were no greater than the typical annual variation, and there were no clear links between diet and breeding allochrony or the differing population trends (more rapid increase in northern than southern giant petrels). The high diet diversity and ability of giant petrels to exploit both carrion and marine prey facilitates coexistence with other sympatric Procellariiformes and explains their favourable population status., Author(s): William F. Mills [sup.1] [sup.2] , Tim I. Morley [sup.1] , Stephen C. Votier [sup.3] , Richard A. Phillips [sup.1] Author Affiliations: (1) grid.478592.5, 0000 0004 0598 3800, British [...]

Prions Pachyptila are the most abundant seabirds in the Southern Ocean and comprise two main groups: those with and without bill lamellae to filter zooplankton. With few exceptions, each breeding location supports at most one species from each of these groups. However, Gough Island supports two morphologically very similar, filter-feeding species: broad-billed P. vittata and MacGillivray's prions P. macgillivrayi. To understand how these two species co-occur in sympatry, we compared the foraging ranges, habitat selectivity, trophic segregation and moult schedules of these species using combined geolocation-immersion loggers. After breeding, both species showed a well-defined westward migration prior to moulting. Moult lasted 11-19 weeks and was significantly longer in MacGillivray's than broad-billed prions. Moulting birds occurred in specific areas within the Argentine Basin, with little overlap between the two species. Habitat analysis revealed species-specific preferences, in particular sea surface temperature. Activity patterns also differed; MacGillivray's prions spent more time in flight, which indicates a more active foraging strategy, relying less on filter feeding. Stable isotope ratios ([delta].sup.15N) in flight feathers were greater in MacGillivray's prion, which is consistent with its less specialized bill morphology resulting in feeding at a higher trophic level. Inter-specific spatial segregation was observed for most of the tracking period, in large part because broad-billed prions breed roughly 3 months earlier than MacGillivray's prions. At Tristan da Cunha, 250 km farther north, where only broad-billed prions breed, they departed, moulted and returned significantly later (15-17 days) than conspecifics from Gough Island, providing evidence for character displacement in sympatry with MacGillivray's prion., Author(s): Christopher W. Jones [sup.1], Richard A. Phillips [sup.2], W. James Grecian [sup.3], Peter G. Ryan [sup.1] Author Affiliations: (1) grid.7836.a, 0000 0004 1937 1151, FitzPatrick Institute of African Ornithology, [...]

Diet analyses can reveal important changes in seabird foraging ecology and, by inference, resource availability and predator-prey dynamics within the wider marine ecosystem. Here, we analysed stomach contents of 1544 grey-headed albatross Thalassarche chrysostoma (GHA) and black-browed albatross T. melanophris (BBA) chicks from Bird Island, South Georgia. We describe dietary shifts (1996-2017), and link those to annual prey availability indices and breeding success. Annual variability in diet was high, and long-term trends in the main components were broadly similar in both albatrosses. Fish consumption (by mass) generally increased over time. Mackerel icefish Champsocephalus gunnari occurrence increased in GHA diets, but was unrelated to local densities derived from fisheries/research cruises. Cephalopod consumption declined until the early 2000s, then plateaued, and the occurrence of the ommastrephid squid Martialia hyadesi declined over time in both albatrosses. In BBAs, Antarctic krill Euphausia superba consumption decreased over time. Conversely, Antarctic krill consumption by GHAs increased until the early 2000s, decreased until the mid-2010s, and increased again in 2017. Antarctic krill consumption was unrelated to local densities based on acoustic surveys, and did not correlate with breeding success. Remotely sensed chlorophyll-a within core foraging areas showed a positive relationship with Antarctic krill in GHA diets, but a negative relationship with M. hyadesi occurrence in both albatross diets. Dietary shifts had consequences for GHA breeding success, which was negatively related to the importance of the cranchiid Galiteuthis glacialis and positively related to M. hyadesi importance. These results highlight the complex mechanisms linking prey availability, diet and breeding success in albatrosses., Author(s): William F. Mills [sup.1] [sup.2], José C. Xavier [sup.1] [sup.3], Stuart Bearhop [sup.2], Yves Cherel [sup.4], Stephen C. Votier [sup.2], Claire M. Waluda [sup.1], Richard A. Phillips [sup.1] Author [...]

Divergent foraging strategies may emerge within a population due to a combination of physiological and environmental factors; yet to persist, neither strategy should offer a consistent selective advantage over the alternative in the long term. Murphy's petrels Pterodroma ultima from Henderson Island (24°20'S, 128°20'W) in the South Pacific Ocean are highly vagile, and exhibit two distinct foraging trip types during incubation; similar proportions of birds undertake either looping trips around the South Pacific Gyre to waters off Peru (hereafter 'East') or trips south-west of the colony towards the Subtropical Front ('South') (mean maximum ranges of c. 3800 or 2000 km from the colony, respectively). However, the relative benefits of the distinct trip types remain unclear. Through tracking birds with GPS and salt-water immersion loggers in 2015, the fine-scale foraging behaviour was examined for East (trip durations: 14.1-19.8 days, maximum ranges 2387-4823 km) and South trips (12.9-25.8 days, 1565-1991 km). Data on behaviour classified from GPS tracks, the number of wet bouts per hour (a proxy for landing rates) and wind speeds, were used to distinguish two distinct foraging modes: birds on East trips spent more time in directed movement, whereas those on South trips spent a greater proportion of time in area-restricted search (ARS) behaviour. East trips were associated with higher overall mass gain, and wet bouts occurred in equal proportions during directed movement and ARS behaviour. This suggests that in unproductive marine environments, it may be more profitable to maximise area covered to increase the chances of encountering prey. Analysis of lower-resolution geolocator data (collected from 2011 to 2014) indicated that individuals were largely consistent in trip type between years. Since birds that conducted East trips were 19% lighter on departure from the colony and experienced more frequent tailwinds on foraging trips, we speculate that these birds may benefit from reduced movement costs, whilst also experiencing reduced competition for foraging opportunities., Author(s): Thomas A. Clay [sup.1] [sup.2] [sup.3] , Steffen Oppel [sup.4] , Jennifer L. Lavers [sup.4] [sup.5] , Richard A. Phillips [sup.1] , M. de L. Brooke [sup.2] Author Affiliations: [...]

The marine ecosystems of the Bering Sea and adjacent southern Chukchi Sea are experiencing rapid changes due to recent reductions in sea ice. Short-tailed shearwaters Puffinus tenuirostris visit this region in huge numbers between the boreal summer and autumn during non-breeding season, and represent one of the dominant top predators. To understand the implications for this species of ongoing environmental change in the Pacific subArctic and Arctic seas, we tracked the migratory movements of 19 and 24 birds in 2010 and 2011, respectively, using light-level geolocators. In both years, tracked birds occupied the western (Okhotsk Sea and Kuril Islands) and eastern (southeast Bering Sea) North Pacific from May to July. In August-September of 2010, but not 2011, a substantial proportion (68 % of the tracked individuals in 2010 compared to 38 % in 2011) moved through the Bering Strait to feed in the Chukchi Sea. Based on the correlation with oceanographic variables, the probability of shearwater occurrence was highest in waters with sea surface temperatures (SSTs) of 8-10°C over shallow depths. Furthermore, shearwaters spent more time flying when SST was warmer than 9°C, suggesting increased search effort for prey. We hypothesized that the northward shift in the distribution of shearwaters may have been related to temperature-driven changes in the abundance of their dominant prey, krill (Euphausiacea), as the timing of krill spawning coincides with the seasonal increase in water temperature. Our results indicate a flexible response of foraging birds to ongoing changes in the sub-Arctic and Arctic ecosystems., Introduction The Bering and southern Chukchi seas are among the most productive marine ecosystems in the world (Highsmith and Coyle 1990; Grebmeier 2012; Hunt et al. 2013). Reductions in sea [...]

Top predators are critical to ecosystem function, exerting a stabilising effect on the food web. Brown skuas are opportunistic predators and scavengers. although skuas are often the dominant land-based predator at seabird colonies, this is the first detailed study of their movements and activity during breeding. The study was carried out at Bird Island, South Georgia (54°00'S, 38°03'W), in the austral summer of 2011/2012 and included GPS data from 33 breeding adults tracked during the late incubation and early chick-rearing periods. Brown skuas spent on average more than 80 % of time in the territory, and it was extremely rare for both partners to leave the territory simultaneously. Much more time was spent foraging at the coast than in penguin colonies and, based on saltwater immersion data, adults never foraged at sea. None of the tracked birds appeared to specialise in catching small petrels at night. Fewer foraging trips were made per day, and hence, more time was spent in the territory, during incubation than chick-rearing. Despite the pronounced sexual size dimorphism, there were no effects of sex on territorial attendance, foraging time or habitat use. Skuas at Bird Island show higher territorial attendance and are less likely to leave the territory unattended than those breeding elsewhere, suggesting closer proximity to more diverse or abundant food resources than at other colonies. The results tie in with previous diet studies, indicating that brown skuas at this site feed mostly on seal placentae and carrion and that birds may rely on a broader range of food resources as the season progresses., Introduction Top predators can exert an important stabilising effect on the food web, keeping prey species in check and preventing the loss of inferior competitors (Paine 1966; Soule et al. [...]

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Miniature geolocator loggers (Global Location Sensing, GLS) that provide daily locations of birds have revolutionised the study of winter ecology and migration patterns of seabirds. A long-term study of ringing recoveries and analyses of heavy metals and pollutants in tissues of Atlantic puffins Fratercula arctica from the Isle of May, south-east Scotland, suggested that this population wintered mainly within the North Sea. However, deployment of GLS devices over the 2007/2008 winter showed that many breeding birds made major excursions into the east Atlantic. This winter was the second of two when survival was extremely low (survival in 2006/2007 and 2007/2008 was 0.696 and 0.695, respectively, compared to the average of 0.922 over the period 1984/1985-2005/2006). These low rates of survival suggested that the unexpected use of the Atlantic might have been associated with unusually poor conditions in the North Sea as indicated by very low breeding success in 2007. Survival rate returned to previous levels in 2008/2009 providing the opportunity to test whether higher survival was associated with birds remaining in the North Sea, or whether movements into the Atlantic are a feature of this population unrelated to survival. Accordingly, geolocators were deployed over the 2009/2010 winter when adult survival was subsequently established to be high (0.913). We found greater support for the hypothesis that winter distribution is not associated with survival. Thus, 8 (40 %) of 20 individuals followed in 2009/2010 went into the Atlantic, a rate not significantly different from 11 (58 %) of the 19 followed in the 2007/2008 winter. Indeed, birds actually spent longer in the Atlantic and used a wider variety of areas in 2009/2010, although the time spent away from the colony was significantly shorter than in 2007/2008. Since our data were from individuals that survived, remaining in or moving out of the North Sea can both be successful strategies during winters when the population as a whole shows either high or low survival rates. Unfortunately, we do not know where birds that died had gone, and hence, the relative survival of birds that did or did not move into the Atlantic. Determining the link between survival and wintering area for any seabird remains a formidable challenge and will have to await the development of technologies that can determine both where and when birds die., Introduction Although the breeding biology of many seabirds is well documented, until recently there was a major gap in our knowledge of seabird migration and wintering areas. What little information [...]

The distribution of Atlantic puffins (Fratercula arctica) from Skellig Michael, south-west Ireland, was investigated using geolocation loggers between the 2010 and 2011 breeding seasons. All tracked birds travelled rapidly west into the North Atlantic at the end of the breeding season in August, with the majority undertaking transatlantic trips from Ireland to the Newfoundland-Labrador shelf. The furthest distance from the colony reached by each bird was not influenced by body mass or sex and was achieved in approximately 20 days. By October, all birds had moved back to the mid Atlantic where they remained resident until returning to the breeding colony. The most parsimonious explanation for the rapid, directed long-distance migration is that birds exploit the seasonally high abundance of prey [e.g., fish species such as capelin (Mallotus villosus) and sandlance (Ammodytes spp.)] off the Canadian coast, which is also utilised by large populations of North American seabirds at this time. Once the availability of this short-term prey resource has diminished, the tracked puffins moved back towards the north-east Atlantic. A relationship between relative abundance of puffins and zooplankton was found in all winter months, but after correcting for spatial autocorrelation, was only significant in November and January. Nevertheless, these results suggest a potential switch in diet from mainly fish during the breeding and early post-breeding periods to zooplankton over the remaining winter period. This study suggests that puffins from south-west Ireland have a long-distance migration strategy that is rare in breeding puffins from the UK and identifies a key non-breeding destination for puffins from Ireland. This has implications for the susceptibility of different breeding populations to the effects of possible climatic or oceanographic change., Introduction Most seabirds undertake long migrations following the breeding season. This behaviour is thought to be an adaptive strategy to avoid seasonal deterioration in climate or increasing scarcity of resources [...]

Sexual segregation in foraging habitat occurs in many marine predators and is usually attributed to competitive exclusion, different parental roles of each sex or niche specialisation associated with sexual size dimorphism. However, relatively few studies have attempted to understand the patterns and underlying drivers of local-scale sexual segregation in marine predators. We studied habitat use, diet and feeding ecology of female and male wandering albatrosses Diomedea exulans, fitted with GPS and stomach-temperature loggers during the chick-rearing period (austral winter) at South Georgia in 2009. During this period, when oceanographic conditions were anomalous and prey availability was low in waters near the breeding colony, the tracked wandering albatrosses showed high consistency in their foraging areas at a large spatial scale, and both males and females targeted sub-Antarctic and subtropical waters. Despite consistency in large-scale habitat use, males and females showed different foraging behaviours in response to oceanographic conditions at a smaller scale. Males appeared to be more opportunistic, scavenging for offal or non-target fish discarded by fishing vessels in less productive, oceanic waters. They exhibited sinuous movements, feeding mostly on large prey and consuming similar amounts of food during the outbound and return parts of the foraging trip. In contrast, females targeted natural productivity hotspots, and fed on a wide variety of fish and cephalopods. They commuted directly to these areas; most prey were ingested on the outbound part of the trip, and they often started their return after ingesting large prey at the farthest point from the colony. Together, these results indicate that sexual segregation in core foraging areas of wandering albatrosses is driven by sex-specific habitat selection due to the low availability of prey in local Antarctic waters. This segregation results in different feeding behaviour at local scales which may be explained by differing breeding roles and degree of parental investment by each sex, with females investing more than males in reproduction. Further investigations are necessary to confirm the existence of this pattern through time under contrasting environmental conditions and to identify the drivers responsible for local-scale sexual segregation in wandering albatrosses., Author(s): Jorge M. Pereira [sup.1] , Vítor H. Paiva [sup.1] , Richard A. Phillips [sup.2] , José C. Xavier [sup.1] [sup.2] Author Affiliations: (Aff1) 0000 0000 9511 4342, grid.8051.c, MARE [...]

The time seabirds have to forage is restricted while breeding, as time at sea must be balanced against the need to take turns with the partner protecting the nest site or offspring, and timing constraints change once the breeding season is over. Combined geolocator-immersion devices were deployed on eleven Imperial Shags (four males and seven females) in Argentina (43°04'S; 64°2'W) in November 2006 and recovered in November 2007. During the breeding season, females foraged throughout the morning, males exclusively in the afternoon, and variability between individuals was low. Outside the breeding season, both sexes foraged throughout the day, and variability between individuals was high. Timing differences may be explained by higher constraints on foraging or greater demands of parental duties experienced by the smaller sex, females in this case. Sexual differences in reproductive role, feeding habits or proficiency can also lead to segregation in timing of foraging, particularly while breeding., Introduction When breeding, seabirds are central-place foragers, forced to adjust their feeding behavior to satisfy the energetic demand of their offspring, as well as their own, under time constraints imposed [...]

There is an urgent need to identify key marine areas for conservation, particularly in the high seas. A range of techniques have been applied to tracking data from higher predators, particularly seabirds and pinnipeds, to determine the areas of greatest use. This study compared three commonly used methods--kernel, first-passage time and state-space modelling--and a new approach, minimum displacement rate, for the analysis of data from the wandering albatross Diomedea exulans of Bird Island, South Georgia, tracked during the chick-rearing period. Applied to a single track, these four models identified similar marine areas as important. The greatest similarity in areas identified occurred when model assumptions were shared (such as slow speed indicating spatial preference) even when methods modelled these assumptions differently (e.g. Bayesian inference versus cumulative density surface). A gridded overlap approach applied to all tracks revealed core areas not apparent from results of any single analysis. The gridded approach also revealed spatial overlap between methods based on different assumptions (e.g. minimum displacement rate and kernel analysis) and between individuals. Although areas identified as important by kernel and first-passage time analysis of a single track were biased towards resting locations during darkness, this does not negate the requirement for their protection. Using the gridded overlap approach, two distinct core regions were identified for the wandering albatross; one close to the breeding colony and another 800 km to the North-West in the high seas. This convenient and pragmatic approach could be applied to large data sets and across species for the identification of a network of candidate marine protected areas in coastal and pelagic waters., Introduction The last two decades have seen a marked increase in research using tracking technologies to understand the ecology and behaviour of animals (Phillips et al. 2007; Aarts et al. [...]

The wandering albatross (Diomedea exulans)is regarded as a generalist predator, but can it be consistent in its foraging niche at an individual level? This study tested short- and long-term consistency in the foraging niche in terms of habitat use, trophic level and, by inference, prey selection. Fieldwork was carried out at Bird Island, South Georgia, in May-October 2009, during the chick-rearing period. Blood (plasma and cells) and feathers for stable isotope analyses (δ[sup.13]C and δ[sup.15]N) were sampled from 35 adults on their return from a foraging trip during which they carried stomach temperature, activity and global positioning system loggers. Results suggest short-term consistency in foraging niche in relation to both oceanic water mass and trophic level, and long-term consistency in use of habitat. Consistent differences between individuals partly reflected sex-specific habitat preferences. The proportion of consistent individuals (i.e., with a narrow foraging niche) was estimated at c. 40 % for short-term habitat and trophic level (prey) preferences and 29 % for longer-term habitat preference, suggesting this is an important characteristic of this population and potentially of pelagic seabirds in general. Foraging consistency was not related to body condition or level of breeding experience; instead, it may reduce intraspecific competition., Introduction According to optimal foraging theory, animals should distribute themselves to maximize their foraging efficiency, which, during breeding, is dependent upon the ability to find food within the range of [...]

Considerable attention has focused on inter- and intraspecific variation in trophic niches of marine predators. Although this has revealed evidence for sexual segregation in distribution in some species, few studies have been able to address sex-related dietary specialisation. Stable isotope analysis of blood cells collected from albatrosses and petrels at South Georgia during chick-rearing indicated a difference in [delta].sup.13C, suggesting that females fed to the north of males, only in two species with male-biased sexual size dimorphism; in no species did sexes differ in trophic level ([delta].sup.15N). Based on a wider review, significant differences between sexes in isotope signatures were much more common in seabirds during the pre-laying or breeding than the nonbreeding period, presumably reflecting greater between-sex partitioning of resources when foraging ranges are more constrained and competition is greater. Sex differences, or their absence, were usually consistent across successive stages during the pre-laying and breeding periods, but not necessarily year-round nor between populations. Significant differences in isotope signatures between males and females were extremely rare in monomorphic species, suggesting a link between sexual size dimorphism and segregation in diet or distribution. Among the Southern Ocean albatrosses, sex differences in [delta].sup.13C suggested the underlying mechanism was related to habitat specialisation, whereas in other size-dimorphic taxa (both male- and female-biased), sex differences were more common in [delta].sup.15N than [delta].sup.13C and therefore more consistent with size-mediated competitive exclusion or dietary specialisation., Author(s): Richard A. Phillips [sup.1] , Rona A. R. McGill [sup.2] , Deborah A. Dawson [sup.3] , Stuart Bearhop [sup.4] Author Affiliations: (1) grid.8682.4, 0000000094781573, British Antarctic Survey, Natural Environment [...]

The long breeding period and high reproductive investment of seabirds make use of resource-rich foraging areas pivotal both during and between breeding seasons. We tracked adult Australasian Gannets (Morus serrator) from their New Zealand breeding colony at Cape Kidnappers to Australia during the non-breeding period to assess wintering behavior and migratory routes for this species. Data from three recovered geolocation sensor (GLS) tags showed that both a male and a female M. serrator, and a hybrid M. capensis x M. serrator migrated across the Tasman Sea to winter in Australian and Tasmanian coastal waters. Tracked birds covered distances of up to 13,000 km on their migration. These movements were consistent with historical records of band recoveries. Received 28 April 2010. Accepted 29 September 2010., Adults of many seabird species, in systems with obligate biparental care, invest heavily during breeding (Brooke 2004). Life history theory predicts that parents must balance the costs of current reproduction [...]

Despite the recent burgeoning in predator tracking studies, few report on seabird activity patterns, despite the potential to provide important insights into foraging ecology and distribution. In the first year-round study for any small petrel, we examined the activity patterns of the white-chinned petrel Procellaria aequinoctialis based on data from combination geolocator-immersion loggers deployed on adults at South Georgia. The petrels were highly nocturnal, flying for greater proportions of darkness than any large procellarid studied so far, except the light-mantled albatross Phoebetria palpebrata. Flight bout durations were short compared with other species, suggesting a dominant foraging mode of small-scale searching within large prey patches. When migrating, birds reduced the proportion of time on the water and increased flight bout duration. Activity patterns changed seasonally: birds flew least during the nonbreeding period, and most frequently during chick-rearing in order to meet higher energy demands associated with provisioning offspring. The degree of their response to moonlight was also stage dependent (greatest in nonbreeding, and weakest in incubating birds), a trait potentially shared by other nocturnal petrels which will have repercussions for feeding success and prey selection. For the white-chinned petrel, which is commonly caught in longline fisheries, these results can be used to identify periods when birds are most susceptible to bycatch, and therefore when use of mitigation and checking for compliance is critical., Author(s): Elizabeth K. Mackley [sup.1] [sup.2] , Richard A. Phillips [sup.1] , Janet R. D. Silk [sup.1] , Ewan D. Wakefield [sup.1] , Vsevolod Afanasyev [sup.1] , Robert W. Furness [...]

Most seabirds die outside the breeding season, but understanding the key factors involved is hampered by limited knowledge of nonbreeding distributions. We used miniature geolocating loggers to examine the movements between breeding seasons of Atlantic puffins Fratercula arctica from a major North Sea colony where numbers have declined in recent years, apparently due to increased overwinter mortality. The most intensively used region was the northwestern North Sea but most puffins also made excursions into the east Atlantic in the early winter. Ringing recoveries previously indicated that adults from British east coast colonies remained within the North Sea and hence were spatially segregated from those breeding on the west throughout the year. Updated analyses of ringing recoveries support results from geolocators suggesting that usage of Atlantic waters is a recent phenomenon. We propose that the increased adult mortality is related to changes in distribution during the nonbreeding period and reflects worsening conditions in the North Sea., Author(s): Michael P. Harris [sup.1] , Francis Daunt [sup.1] , Mark Newell [sup.1] , Richard A. Phillips [sup.2] , Sarah Wanless [sup.1] Author Affiliations: (1) grid.8682.4, 0000000094781573, Centre for Ecology [...]

The study of long-distance migration provides insights into the habits and performance of organisms at the limit of their physical abilities. The Arctic tern Sterna paradisaea is the epitome of such behavior; despite its small size ( at-sea hotspot | global wind systems | geolocator | trans-equatorial migration doi/10.1073/pnas.0909493107

To link to full-text access for this article, visit this link: http://dx.doi.org/10.1016/j.anbehav.2009.09.029 Byline: Jose P. Granadeiro (a), Hany Alonso (b), Vitor Almada (b), Dilia Menezes (c), Richard A. Phillips (d), Paulo Catry (b) Abstract: Several species of seabirds show cyclic patterns of attendance at their nesting colonies. We examined the patterns of variation in the numbers of Cory's shearwater at three colonies (two oceanic and one located on the continental shelf), including the world's largest, at Selvagem Grande, Madeira, Portugal and considered several hypotheses concerning their causal mechanisms. At Selvagem Grande, cycles were exceptionally marked and regular, with a periodicity ranging from 7.8 to 11 days, and involved both breeders and nonbreeders. In contrast, variation in numbers was aperiodic at a nearby and much smaller colony (Selvagem Pequena), and also at the colony located off the Portuguese coast (Berlenga Island). We found no relationships between number of birds ashore and environmental variables such as wind direction and speed or lunar cycle. Cycles did not seem to be driven by oscillations in food availability or accessibility, given that they did not correlate with daily chick growth rates (which were acyclic) or diet. Despite their regularity, cycles were slightly out of phase in different sectors of Selvagem Grande, which suggests that social interactions at the colony could act as an entrainment agent for an endogenous rhythm, and so cycles are probably more likely to occur in large and dense colonies. Observations are consistent with the hypothesis that cycles facilitate social interactions by maximizing the probability of encounters at the colony. However, the exact mechanisms through which these remarkable cycles are controlled are still completely unknown, and clearly further research is needed. Author Affiliation: (a) Centro de Biologia Ambiental, Museu Nacional de Historia Natural, Universidade de Lisboa, Portugal (b) Eco-Ethology Research Unit, Instituto Superior de Psicologia Aplicada, Portugal (c) Parque Natural da Madeira, Portugal (d) British Antarctic Survey, Natural Environment Research Council, U.K. Article History: Received 25 June 2009; Revised 30 July 2009; Accepted 17 September 2009 Article Note: (miscellaneous) MS. number: 09-00427

By extracting energy from the highly dynamic wind and wave fields that typify pelagic habitats, albatrosses are able to proceed almost exclusively by gliding flight. Although energetic costs of gliding are low, enabling breeding albatrosses to forage hundreds to thousands of kilometers from their colonies, these and time costs vary with relative wind direction. This causes albatrosses in some areas to route provisioning trips to avoid headwind flight, potentially limiting habitat accessibility during the breeding season. In addition, because female albatrosses have lower wing loadings than males, it has been argued that they are better adapted to flight in light winds, leading to sexual segregation of foraging areas. We used satellite telemetry and immersion logger data to quantify the effects of relative wind speed, sex, breeding stage, and trip stage on the ground speeds ([V.sub.g]) of four species of Southern Ocean albatrosses breeding at South Georgia. [V.sub.g] was linearly related to the wind speed component in the direction of flight ([V.sub.wf]), its effect being greatest on Wandering Albatrosses Diomedea exulans, followed by Black-browed Albatrosses Thalassarche melanophrys, Light-mantled Sooty Albatrosses Phoebatria palpebrata, and Gray-headed Albatrosses T. chrysostoma. Ground speeds at [V.sub.wf] = 0 were similar to airspeeds predicted by aerodynamic theory and were higher in males than in females. However, we found no evidence that this led to sexual segregation, as males and females experienced comparable wind speeds during foraging trips. Black-browed, Gray-headed, and Light-mantled Sooty Albatrosses did not engage in direct, uninterrupted bouts of flight on moonless nights, but Wandering Albatrosses attained comparable [V.sub.g] night and day, regardless of lunar phase. Relative flight direction was more important in determining [V.sub.g] than absolute wind speed. When birds were less constrained in the middle stage of foraging trips, all species flew predominantly across the wind. However, in some instances, commuting birds encountered headwinds during outward trips and tail winds on their return, with the result that [V.sub.g] was 1.0-3.4 m/s faster during return trips. This, we hypothesize, could result from constraints imposed by the location of prey resources relative to the colony at South Georgia or could represent an energy optimization strategy. Key words: albatrosses; ARGOS errors; central-place foraging; dynamic or gust soaring; flight speed; habitat preference; niche specialization; platform terminal transmitters, PTT; satellite tracking; seabird; sexual segregation; South Georgia.

To authenticate to the full-text of this article, please visit this link: http://dx.doi.org/10.1111/j.1467-8551.2008.00564.x Byline: Julie Froud (*), Sukhdev Johal (*), Adam Leaver (*), Richard Phillips (*), Karel Williams (*) Abstract: This paper aims to explain the recent defensive strategy of downsizing at the BBC. The paper rejects the analysis of both industry practitioner and neoclassical economics academic critics who represent the corporation as an all-powerful, abusive player in a market and instead develops an alternative concept of the business model, which focuses the pressures of financial viability and stakeholder credibility to explain the restructuring. We argue that the BBC's business model is stressed because it struggles to deliver what key stakeholders want and expect from the corporation, from a pot of revenue that is limited by regulation. The BBC's problem is compounded by demands for more programming hours following its move into digital and by the increasingly formalized demands of regulators on behalf of an absent consumer. The paper concludes that without reflexive, business-model-centred regulation, it is likely that the BBC's business model will become unsustainable. Author Affiliation: (*)Centre for Research on Socio Cultural Change, 178 Waterloo Place, University of Manchester, Oxford Road, Manchester M13 9PL, UK Article note: Corresponding author email: adam.leaver@mbs.ac.uk

In his article for this issue of Economy and Society, Narayanan (2008) presents a critique of our earlier work on Malaysian industrialization (Henderson & Phillips 2007). Narayanan's objections are entirely empirical and they take the form of' suggesting that had we looked at the 'right' data or 'properly' interpreted the data we provide, we would have seen that our hypothesis that Malaysia's industrialization project was 'stalling' was either wrong, or, if plausible, was unproven. In this response, we show that Narayanan is largely mistaken in his critique. While his contribution contains errors of interpretation, we argue that he is mistaken predominantly because he has misunderstood the methodological basis of our analysis. Specifically, he has failed to grasp our theoretical object and thus has been unable to identify the appropriate empirical terrain for assessing its significance and condition. Keywords: Malaysia; industrialization; foreign direct investment; methodology.

Enhanced levels of microbial biomass and activity in the rhizosphere arise from labile C released from roots but the factors that mediate such 'rhizosphere effects' are poorly understood. We hypothesized that the magnitude of rhizosphere effects would be reduced by increases in soil fertility, consistent with plant C allocation theory, which predicts decreased C flux to roots in fertile soils. Replicate plots of sugar maple (Acer saccharum Marshall) and northern red oak (Quercus rubra L.) at the Turkey Hill Plantations, New York, and yellow birch (Betula allenghaniensis Britton) at the Hubbard Brook Experimental Forest, New Hampshire, were fertilized from 2001 to 2003. In fall of 2003, rhizosphere and bulk soils were sampled from the organic horizon of control and fertilized plots and analyzed for microbial biomass and nutrient transformation rates. In general, fertilization reduced microbial biomass and activity in plots of all three species, and the magnitude of such effects was generally greater in rhizosphere than in bulk soil. In red oak soils, fertilization reduced rhizosphere effects on microbial biomass, net N mineralization rates, and phosphatase enzyme activity (P = 0.026, 0.091, and 0.061, respectively). In contrast, fertilization only reduced rhizosphere effects on microbial biomass in sugar maple soils (P = 0.019). In yellow birch soils, fertilization decreased rhizosphere effects on microbial biomass (P = 0.015). These results suggest that soil fertility may mediate the degree to which roots affect microbial activity in forest soils, possibly due to fertilizer-induced shifts in belowground C supply. Abbreviations: DWE, dry weight equivalent; HBEE Hubbard Brook Experimental Forest; MB, microbial biomass; MRA, maximum respiratory activity; THE Turkey Hill Plantations

The study of animal foraging behaviour is of practical ecological importance (1), and exemplifies the wider scientific problem of optimizing search strategies (2). Levy flights are random walks, the step [...]

Evans and Rauch (1999) have demonstrated that there is a strong relationship between the 'Weberianness' of a national state's bureaucratic capacities and its record on economic growth. Using their data set and the best available data sets on the incidence of income poverty, this article investigates whether there is a relationship between a Weberian notion of effective bureaucratic capacities and the state's ability to reduce poverty. Using the same time frame as Evans and Rauch (1970-90) and analysing data for 29 developing and middle income countries, the article concludes that, in general, there does indeed seem to be a relationship between states with effective 'Weberian' public institutions and their ability to reduce poverty. KEY WORDS bureaucracies / economic growth / poverty / states

The Karakoram fault is a major NW-SE-aligned dextral strike-slip fault that has accommodated minor extrusion of thickened Tibetan crust since the Mid-Miocene. Controversy exists as to whether the fault is lithospheric in scale, resulting in synkinematic metamorphism and granite melting, or whether it is an upper crustal fault cutting through earlier metamorphic and granitic rocks. Structural, metamorphic and geochronological data from the Karakoram fault show that: (1) metamorphism and granite genesis occurred prior to strike-slip shearing, meaning that U-Pb zircon and monazite ages from deformed granites cannot be used as proxies for dating strike-slip shearing, only as a maximum age constraint on initiation of ductile shearing; (2) sheared Miocene leucogranites along the fault are equivalent to the Baltoro granite batholith, offset 40-150 km along the fault, and show a strain gradient from high ductile strain to undeformed away from the fault; (3) [sup.40]Ar/[sup.39]Ar and fission-track ages of K2 orthogneisses and Gasherbrum diorite show that rocks adjacent to the fault cooled during Cretaceous to mid-Tertiary time, and were not reset during shearing, indicating that shear heating was not important; (4) the Gasherbrum diorite, adjacent to the fault, has never been at depths greater than 6 km since the early Cretaceous and less than 3 km of material has been eroded off since then. We conclude that the Karakoram fault has had no significant influence on regional metamorphism or melting, at least at levels exposed today, and is restricted to the upper-middle crust.

This paper examines the relation of particular forms of social and labour market policy to economic development. Taking the history of Malaysian industrialization as its empirical case, the paper assesses the unintended consequences of redistribution policy, on the one hand, and migration policy, on the other, for the limited upgrading of the country's electronics industry. It argues that, while the former has been central to social harmony in Malaysia's multi-racial society, it has contributed to the underdevelopment of small and medium-sized firms capable of linking with the TNCs on the basis of knowledge-intensive and higher value-added operations. Migration policy, on the other hand, has allowed manufacturers to have continued access to supplies of low-cost, lower-skilled labour that have released the pressures that would otherwise have been there for technological and skill upgrading in the electronics industry. Only in Penang, where regional state institutions have intervened to encourage SME upgrading, has the national picture been moderated. Malaysia's industrialization project emerged at time when export competition in manufactured commodities was less intense than it is now. Largely as a result of federal government priorities and for other reasons explored in the paper, advantage was not taken of this 'window of opportunity'. As a consequence, the country's industrialization project--exemplified by its electronics industry--is now 'stalling' in the sense that it remains locked into low- to medium-technology operations. With the rise of China as a manufacturing exporter, this is a dangerous situation for a country's principal industry to be in. Keywords: social policy; migration; labour markets; state institutions; economic development; industrialization; Malaysia.

This paper takes issue with arguments emanating from the global social policy literature that neoliberal policy agendas have been largely a consequence of the interplay of international agencies with indigenous reform interests. While relevant, such arguments grasp only part of the story of social policy change. By means of a case study of Hungary between 1990 and 2002, this article emphasises the role played by the bureaucratic reconstitution of the state and changing forms of national economic governance in the explanation of social policy change. We show how the bureaucratic redesign of the Hungarian state generated a 'finance-driven' form of economic governance with the state bureaucracy reconfigured around the fiscal control of the Finance Ministry. These changes had significant implications, not simply for social expenditure, but for the intellectual nature and bureaucratic space for social policy-making. Whereas critiques of neoliberal social policy reform tend to focus on the ideological nature of the projects, this analysis highlights the need to develop visions of, and arguments for, an alternative to the finance-driven forms of economic governance that have become the de facto bureaucratic archetype for re-designing welfare states.

To link to full-text access for this article, visit this link: http://dx.doi.org/10.1016/j.anbehav.2005.10.030 Byline: Paulo Catry, Richard A. Phillips, Jaume Forcada, John P. Croxall Abstract: With rare exceptions, avian offspring are continuously attended by one parent for at least the first few days after hatching. The duration of this phase of the nesting cycle is regulated by a trade-off between the benefits of brooding/guarding and those of foraging. We manipulated offspring age in grey-headed albatrosses, Thalassarche chrysostoma, by swapping chicks between nests. Parents given chicks 6 days older than their own shortened brooding by only 1.2 days, and parents given chicks 6 days younger than their own extended brooding by 1.4 days. Despite being relatively unresponsive to chick age, parents adjusted brooding in relation to calendar date and to chick condition. The results suggest that adults do not use chick age per se as a cue, but instead probably use an internal timer, and fine-tune the decision to end brooding according to date and chick mass. The duration of brood guarding did not correlate with adult body condition, suggesting that adults had a sufficient safety margin to allow them to respond to chick needs without compromising their own residual reproductive value. Chick survival at the end of brood guarding was strongly dependent on calendar date (early and late chicks suffered higher mortality), which suggests that grey-headed albatrosses benefit from breeding synchronously. We conclude that the length of the brood-guarding period is dependent on chick condition and seasonal variation in chick predation risk. Author Affiliation: (a) British Antarctic Survey, Natural Environment Research Council (a ) Unidade de Investigacao em Eco-Etologia, Instituto Superior de Psicologia Aplicada Article History: Received 24 February 2005; Revised 7 June 2005; Accepted 20 October 2005 Article Note: (miscellaneous) MS. number: 8479

A study on relationship between length of brooding by grey-headed albatrosses, Thalassarche chrysostoma, and chick age is analyzed.

Abstract. Previous research on the effects of tree species on soil processes has focused primarily on the role of leaf litter inputs. We quantified the extent to which arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) tree species influence soil microbial activity and nutrient availability through rhizosphere effects. Rhizosphere soil, bulk soil, and fine roots were collected from 12 monospecifc plots (six AM and six ECM tree species) planted on a common soil at the Turkey Hill Plantations in Dryden, New York. Rhizosphere effects were estimated by the percentage difference between rhizosphere and bulk soil samples for several assays. Rhizosphere effects on soil microbes and their activities were significant for ECM species but in only a few cases for AM species. In AM tree species, microbial biomass, net N mineralization, and phosphatase enzyme activity in the rhizosphere were 10-12% greater than in bulk soil. In ECM tree species, rhizosphere effects for microbial biomass, C mineralization rates, net N mineralization, and phosphatase activity were 25-30% greater than bulk soil, and significantly greater than AM rhizosphere effects. The magnitude of rhizosphere effects was negatively correlated with the degree of mycorrhizal colonization in AM tree species (r = -0.83) and with fine root biomass (r = -0.88) in ECM tree species, suggesting that different factors influence rhizosphere effects in tree species forming different mycorrhizal associations. Rhizosphere effects on net N mineralization and phosphatase activity were also much greater in soils with pH < 4.3 for both AM and ECM tree species, suggesting that soil pH and its relation to nutrient availability may also influence the magnitude of rhizosphere effects. Our results support the idea that tree roots stimulate nutrient availability in the rhizosphere, and that systematic differences between AM and ECM may result in distinctive rhizosphere effects for C, N, and P cycling between AM and ECM tree species. Key words: arbuscular mycorrhizae, ectomycorrhizae; rhizodeposition; rhizosphere C flux; root exudates; tree species effects.

Many birds show a surprising degree of intraspecific variability in migratory tendency and choice of wintering site. In this study, we tracked the seasonal movements of 35 nonbreeding Black-browed Albatrosses Thalassarche melanophrys from South Georgia, including 24 birds followed in two consecutive years. This revealed consistent patterns of status-related, sex-specific, and individual variation in wintering strategies, and provided the first description of the summer distribution of failed/deferring breeders. Individuals exhibited a striking degree of site fidelity, returning to the same region (southwest Africa or Australia) and showing correlated centers of distribution, as well as remarkable consistency in the chronology of their movements, in consecutive years. Nonetheless, a degree of behavioral flexibility remained, and particularly on the return migration, birds moved between, or bypassed, alternative intermediate staging sites depending on local circumstances. Initiation of the outward migration varied according to breeding status, timing of failure, and sex: deferring breeders and those that failed early departed two months before successful birds, and successful females departed 1-2 weeks earlier than males. Sex-related latitudinal variation in distribution was also apparent, with females wintering farther north within the Benguela system. Moreover, the only migrant to Australia was a male, supporting an apparent tendency for male-biased breeding dispersal inferred from genetic analyses. Distribution and timing of movements appeared in general to relate to avoidance of competition from congeners and conspecifics from other populations. From a conservation perspective, the study indicated that, for the declining Black-browed Albatross population at South Georgia, the primary focus should be toward improving the management (especially reducing bycatch levels) of fisheries in the central and eastern South Atlantic. Key words: conservation; differential migration; interspecific competition; sexual segregation; winter philopatry.

UNPACKING AND APPLYING TO THE IMPERIAL FIELD Michel Foucault's analysis of sexuality as 'an especially dense transfer point for relations of power,' historians of sexuality and postcolunial critics have identified [...]

This paper analyzes the factors affecting the conditional probability that defaulted residential mortgage loans will foreclose. We analyze a large national sample of conventional loans, which have been in default [...]

A Gray-headed Albatross (Thalassarche chrysostoma) was recorded traveling, in the course of a foraging trip, at a minimum average ground speed of >110 km [h.sup.-1] for ~9 h with virtually no rest. After taking into account the sinuosity of albatross flight, actual mean ground speed was predicted to be [greater than or equal to] 127 km [h.sup.-1], achieved in association with high tailwinds during an Antarctic storm. Despite its high speed and the storminess of the sea, the albatross still managed to successfully locate and capture prey at a rate comparable to that achieved under less extreme conditions. This individual's performance suggests that albatrosses have the capacity to maintain positive energy budgets while quickly covering long distances and taking advantage of the strong winds that are frequent in the Southern Ocean. Received 3 November 2003, accepted 13 May 2004.

Author(s): Stephen C. Votier (corresponding author) [1]; Robert W. Furness (corresponding author) [1]; Stuart Bearhop [1, 2]; Jonathan E. Crane [1]; Richard W. G. Caldow [3]; Paulo Catry [4]; Kenny [...]

The effectiveness of catastrophic-loss index options in hedging hurricane losses for Florida insurers is analyzed, using a windstorm simulation model developed by Applied Insurance Research.

Effects of deployment of miniaturized transmitters and loggers have been well studied in penguins, but much less so in flying seabirds. We examined the effects of satellite tag (platform terminal transmitter, PTT) deployment in Black-browed (Thalassarche melanophris) and Gray-headed (T. chrysostoma) albatrosses at South Georgia and reviewed the recent literature for other albatrosses and petrels. In our study, although a few individuals may have slightly extended their foraging trips, overall there was no significant difference in trip duration, meal mass, breeding success, or rate of return in the next season between birds with PTTs and controls. By comparison, most other studies of albatrosses and petrels recorded extended trip durations and, in some cases, high rates of nest desertion following PTT attachment. That occurred particularly where transmitter loads exceeded 3% of adult mass. Extended trip durations may result from reduced flight efficiency, as well as the effect of capture and temporary restraint, but affected birds seem nonetheless to commute to representative foraging areas. To minimize device effects, we suggest that transmitter loads be reduced to a minimum, use of harnesses be avoided (particularly for breeding season deployments when tape attachment to feathers is an effective alternative), and careful attention be given to limiting handling times during incubation when some species are particularly sensitive to disturbance. Received 17 October 2002, accepted 14 June 2003.

To the Editor: Buruli ulcer, a necrotizing skin disease caused by Mycobacterium ulcerans, is highly endemic to West Africa (1,2) and is characterized by large ulcerations on the lower limbs [...]

Cooperative breeding is unusual in seabirds, but recorded in varying proportions in Brown Skuas (Stercorarius [= Catharacta] lonnbergi). With no evidence for kin selection or delayed dispersal, nor for any reproductive advantage, the reason for its occurrence in skuas is uncertain. This study describes the incidence, composition, and breeding characteristics of trios of Brown Skuas at Bird Island, South Georgia. Trios were recorded in nine seasons, including a polygynous and a polyandrous trio found in both 2000-2001 and 2001-2002. In some years, birds hatched three chicks from supranormal clutches, which is exceptional in skuas and jaegers, yet their overall performance was no better than monogamous pairs. I suggest that compared with closely related taxa, Brown Skuas have a slight behavioral or genetic predisposition that facilitates cooperative breeding. Cooperation conveys a minor advantage in territory acquisition, but none in terms of reproductive success (particularly for polygynous groups), and consequently has not proliferated at most colonies. Key words: cooperative breeding, habitat saturation, polygyny, supranormal clutch.

Numerous explanations exist for the evolution of reversed size dimorphism in raptorial species. A recent study concluded that reversed size dimorphism in skuas and jaegers was probably not attributable to breeding-role specialization, but that there was evidence for sexual selection, and in particular intrasexual competition by females for males. Our study tested the applicability of those conclusions for Southern (or Brown) Skuas (Stercorarius skua lonnbergi) breeding in South Georgia. Clutch volume was related positively to size and condition of females and negatively to condition of males, but there was no evidence of assortative mating for size or condition within pairs. Potential explanations for the discrepancy between this and previous studies are that size is less closely correlated with individual quality because of highly diverse foraging strategies, territory quality is a confounding factor, or because lower aggression in Southern Skuas reduces the necessity for small females to avoid large males.