Colony Attendance Research Articles

Little auks under the midnight sun: diel activity rhythm of a small diving seabird during the Arctic summer

Many animal species exhibit a diel, 24-hr pattern of activity, which is steered by timing cues, with the daily light–dark cycle considered the most powerful. This cue, however, is reduced in polar zones under continuous daylight conditions associated with the midnight sun. The rhythm of animal behaviour under such conditions is poorly understood. Here, we examine periodicity and patterns of daily activity (colony attendance and foraging) in a High-Arctic seabird, the little auk (Alle alle). We demonstrated a regular rhythm of colony attendance at the population level, with birds being the most abundant in the colony during hours of relatively low sun elevation. This pattern is likely to be associated with predation pressure that may be perceived by birds as lower during hours with low sun elevation, because of better predator detectability. Regarding rhythms at an individual level, however, we found the most common periodicity to be 23.2 hr (range from 19.9 hr to 30.8 hr) but no clear pattern of daily colony attendance of individuals. Such a flexibility in daily rhythms indicates that individuals may become arrhythmic in regard to the 24-hr environmental cycle, despite regularities observed at the population level. Finally, we compared males and females in terms of daily activity patterns but we did not find significant sex differences.

Flying to the moon: Lunar cycle influences trip duration and nocturnal foraging behavior of the wedge-tailed shearwater Ardenna pacifica

Lunar phase and illumination are known to affect nocturnal behavior of many organisms, particularly through predator-prey interactions. Visual predators can benefit from higher light levels to increase their activity, while prey may decrease their activity to avoid predation. The lower number of nocturnal seabirds observed on colonies during full moon nights has been mostly interpreted as a predation avoidance strategy. However, it is also possible that shearwaters take advantage of the moon's illumination to feed also at night, and stay at sea to forage during full moon nights. We used miniaturized GPS-loggers to obtain 179 tracks from 99 wedge-tailed shearwaters breeding in New Caledonia, to investigate moonlight effects on individual behavior. Lunar phase significantly predicted self-provisioning trip duration, with individuals performing longer trips around the full moon. However, this relationship was not significant during chick-provisioning trips when adults have to frequently return to the colony. Adults mostly returned to the colony during moonlit periods, refuting the predation avoidance theory. Tracked individuals showed an unexpectedly high amount of nocturnal foraging activity (28% of total activity), positively influenced by the presence of the moon. δ15N stable isotope values were significantly related to the percentage of nocturnal foraging, but with a weak relationship, impeding our ability to confirm that wedge-tailed shearwaters fed on different prey when foraging at night. This study suggests that reduced colony attendance around the full moon may be linked to greater at-sea foraging opportunities in distant oceanic areas than to increased predation risk on land.

Monitoring Surveys of the Parakeet Auklet (Aethia psittacula, Charadriiformes, Alcidae): Daily and Seasonal Attendance Patterns in North Pacific Colonies

The daily and seasonal patterns of colony attendance for the parakeet auklet were examined on Buldir and St. Paul islands, where fieldwork was conducted in 2001, 2002, and 2004, respectively. The timing of breeding events for the parakeet auklet was measured by monitoring the productivity plots on Buldir Island, and the timing of chick hatching was determined by observing birds that carried food to chicks on St. Paul Island. On Buldir Island, direct observations of birds on water were used to collect data on parakeet auklet attendance as it was impossible to count the birds directly on land because of lush vegetation covering the slope of the colony. The birds were counted on a designated plot on water below the colony. On St. Paul Island, observations of birds on both the water and the cliffs were used to collect data on parakeet auklet attendance patterns. On each count day, the birds present at the plot were counted every 15 min. Parakeet auklets have two peaks in their attendance pattern on the water near the colony on Buldir Island. As the breeding season progressed, the peak timing of birds moved slightly toward the afternoon. On St. Paul Island, prior to egg-laying, the numbers of parakeet auklets attending the breeding colony peaked in the morning. As the breeding season progressed, the peak of bird attendance at the colony broadened and shifted toward the evening. During the chick-rearing period, birds arrived at the land plot throughout the day, but there was a peak in the numbers attending in the evening. The observations showed that the weather conditions strongly influenced the numbers of birds present at the plot. Strong winds decreased the number of parakeet auklets in the nearshore area. On windy days, the birds left the plot earlier than usual.

Behavioural plasticity in the early breeding season of pelagic seabirds - a case study of thin-billed prions from two oceans

BackgroundIn long-lived seabirds that migrate large distances independently of each other, the early part of the breeding season is crucially important for a successful reproductive attempt. During this phase, pair bonds are re-established and partners coordinate their breeding duties. We studied the early breeding season in Thin-billed prions Pachyptila belcheri breeding in the Atlantic Ocean (Falkland/Malvinas Islands) and Indian Ocean (Kerguelen). Despite overlap in the wintering areas, these two populations exhibit differences in their timing and direction of migration. We hypothesised that these differences would influence behaviour during the early breeding season.ResultsIn line with our hypothesis, we found very strong differences in colony attendance patterns. Thin-billed prions of the Falkland population spent the late winter period over shelf waters close to the colony, first arrived back at the colony in September, and attended the nests interruptedly for one month, before departing on a pre-laying exodus. In contrast, Kerguelen birds remained in the non-breeding areas until mid-October and spent much less time attending the burrow before their pre-laying exodus. Despite this asynchronous arrival to the two colonies, the subsequent patterns resulted in remarkably synchronous incubation in both populations, with males taking on the first long incubation shift in late November, whereas females returned to sea soon after egg laying. During the pre-laying exodus and incubation, Thin-billed prions from the Falklands spread north over the Patagonian Shelf, while prions from Kerguelen travelled much further, reaching southern oceanic waters and moved at faster speeds (> 400 km per day). Although prions from Kerguelen moved much further, their isotopic niches were considerably narrower, suggesting a stronger dependence on Antarctic waters.ConclusionsThe study thus suggests that Thin-billed prions show a high intraspecific plasticity in their use of either neritic or oceanic waters during the early breeding season. Breeding birds from the Falkland Islands can exploit an extensive shelf area, while Kerguelen birds have adapted to the need to forage in distant southern open waters. This difference in foraging ecology may thus have shaped the phenology of the early breeding phase.

Behavioral responses of Australian fur seals to boat approaches at a breeding colony

In Australia, a multi-million-dollar industry is based on viewing the Australian fur seal (Arctocephaluspusillusdoriferus), predominantly through boat visits to breeding colonies. Regulation of boat approaches varies by site and no systematic investigations have been performed to inform management guidelines. To investigate possible effects of disturbance, experimental boat approaches were made to a colony at Kanowna Island in northern Bass Strait and seal responses were monitored using instantaneous scan sampling. Colony attendance (individuals remaining ashore) was found to be influenced by approach distance and time of day, but was not affected by environmental variables or season, whereas onshore resting behavior was influenced by approach distance, time of day, ambient temperature and wind direction. Onshore resting behavior decreased following experimental boat approaches to 75 m, but changes in abundance of individuals ashore were not observed at this distance. In contrast, approaches to 25 m elicited a strong response, with a steep decline in the number of individuals ashore. This response was strongest when approaches occurred in the morning, with a decline of approximately 47% of individuals, compared to a decline of 21% during afternoon approaches. With regard to onshore resting behavior, afternoon approaches to 75 m led to minimal response. The remaining three combinations of approach distance and time of day had a similar pattern of reductions in the proportion of individuals engaging in onshore resting behavior. The strongest response was again seen during approaches to 25 m conducted in the morning. These behavior changes suggest that unrestricted boat-based ecotourism at Australian fur seal colonies has the potential to increase energy expenditure and reduce the number of seals ashore. Increasing minimum approach distances to ≥75 m and/or restricting visits to afternoons may minimize these impacts at Kanowna Island during the post-molt and non-breeding seasons. As several studies have demonstrated considerable intra-species variation in seal responses to boat approaches, research at other colonies is needed before these findings can be generalized to the remainder of the Australian fur seal population.

Year-round colony attendance patterns for the Gentoo Penguin (Pygoscelis papua) at Martillo Island, Tierra del Fuego, Argentina

ABSTRACT Long-term monitoring of seabird breeding cycles and breeding success is essential for conservation and a valuable indicator of ecological change, but is logistically expensive and time consuming. We describe the annual cycle and year-round colony attendance patterns for Gentoo Penguins Pygoscelis papua at Martillo Island, Beagle Channel, by analyzing camera trap photographs. We deployed a Reconyx HC500 Hyperfire trail camera programmed in time-lapse mode to take single pictures once per hour year-round (27 Feb 2013 until 27 Feb 2014). The nonreproductive stage began 23 April 2013 when adults completed molt and ended 24 September 2013 when the first pair formation was observed. In general Gentoos Penguins performed daily trips, and the number of adults present at 1900 h ART was not significantly different at any stage except the late chick-rearing and premolt stages. Breeding success was 0.92 chicks per nest. Small variations in day length had a significant influence on penguin's trip duration dur...

Penguin colony attendance under artificial lights for ecotourism

Wildlife watching is an emerging ecotourism activity around the world. In Australia and New Zealand, night viewing of little penguins attracts hundreds of thousands of visitors per year. As penguins start coming ashore after sunset, artificial lighting is essential to allow visitors to view them in the dark. This alteration of the nightscape warrants investigation for any potential effects of artificial lighting on penguin behavior. We experimentally tested how penguins respond to different light wavelengths (colors) and intensities to examine effects on the colony attendance behavior at two sites on Phillip Island, Australia. At one site, nocturnal artificial illumination has been used for penguin viewing for decades, whereas at the other site, the only light is from the natural night sky. Light intensity did not affect colony attendance behaviors of penguins at the artificially lit site, probably due to penguin habituation to lights. At the not previously lit site, penguins preferred lit paths over dark paths to reach their nests. Thus, artificial light might enhance penguin vision at night and consequently it might reduce predation risk and energetic costs of locomotion through obstacle and path detection. Although penguins are faithful to their path, they can be drawn to artificial lights at small spatial scale, so light pollution could attract penguins to undesirable lit areas. When artificial lighting is required, we recommend keeping lighting as dim and time-restricted as possible to mitigate any negative effects on the behavior of penguins and their natural habitat.

Population differentiation in whiskered aukletsAethia pygmaea: do diurnal and nocturnal colonies differ in genetics, morphometry and acoustics?

Despite a great number of empirical studies, the mechanisms of population differentiation and the factors that influence this process, particularly in seabirds, remain insufficiently understood. Here we analyzed population structure in the whiskered auklet Aethia pygmaea, a previously poorly studied alcid species with unusual differentiation in colony attendance rhythms (i.e. diurnal in the Sea of Okhotsk vs nocturnal in the Bering Sea), and examined the influence of it on intraspecific differentiation. For this study, we analyzed morphometric measurements, acoustic variables, mitochondrial control region fragment and five microsatellite loci from nine whiskered auklet colonies spanning the breeding range. Previous research has shown a clinal morphometric variation in this species. We build on this analysis by adding auklets from more colonies, for the first time analyzing vocalizations from different colonies and genetic structure of this species. Our data supports a clinal variation in morphometric and acoustic characters with the largest size and the lowest call frequency in western birds, and the smallest size and highest call frequency in the east. We also found two mitochondrial lineages – whiskered auklets from colonies in the Sea of Okhotsk and the Commander Is. (Bering Sea) and from the Aleutian Is. (Bering Sea), that were presumably formed during Sangamonian interglacial period (115 000–130 000 years ago). Genetic clusters found did not reflect differences in colony attendance rhythms, suggesting that they were shaped by other factors (e.g. differences in predator pressure) and are unlikely to have participated in the formation of population structure. Colony fidelity, mobility of birds, proximity of foraging grounds and location of colonies in relation to seasonal ice pack, seem to be more likely determinants of population differentiation.

Fitting in with the crowd: the role of prospecting in seabird behavioural trends

Prebreeding, subadult seabirds have been documented prospecting or visiting multiple sites throughout the breeding season to gather information on colony reproductive success, identify suitable habitat, evaluate prey abundance, and locate potential partners; however, many aspects of prospector biology remain unknown. We explored prospector behaviour as a means of furthering our understanding of postnatal seabird dispersal and colony attendance using Least Auklets (Aethia pusilla (Pallas, 1811)) and Crested Auklets (Aethia cristatella (Pallas, 1769)) breeding at Gareloi Island, Alaska, in 2014 and 2015. We recorded age class, length of time spent on the colony, and behaviour for individuals attending a study plot over the course of two breeding seasons. Although prospectors typically spent more time on the colony surface than adults, prospectors rarely socialized with conspecifics during their visits to the colony, possibly due to the absence of a citrus-like feather odour used in olfactory communication. Additionally, we found substantial differences between observed and predicted data between years, demonstrating that other factors (likely prey abundance or quality) influenced behaviour in 2015. Our results suggest that the collective knowledge of seabird prospecting behaviour is not necessarily transferable between taxa and there may be a range of strategies employed by prospectors when assessing colonies.

Nocturnal colony attendance by Common Guillemots Uria aalge at colony in Shetland during the pre-breeding season

Time-lapse photography was used to describe daily and seasonal trends in attendance by Common Guillemot Uria aalge at a colony in Shetland prior to the breeding season, including detection of nocturnal presence. A camera took a photo every 30 minutes from 30 January until 21 April 2015. A total of 3,435 photos were analysed, of which 3,232 photos allowed birds to either be accurately counted (2,552) or estimated (680) within a representative plot. High quality moonlit shots showed that large numbers of Common Guillemots were present ashore at Sumburgh throughout the night, while manipulated non-moonlight night photos revealed attendance at the colony, even when counting was not possible. Clear cycles of attendance at the colony were apparent with day-time peaks of 90–120 birds occurring on average every 7 days over the study period. Pre-breeding attendance is described, as is the nocturnal presence of Common Guillemots in the three months prior to breeding.

The need for year-specific correction factors (k values) when converting counts of individual Common Guillemots Uria aalge to breeding pairs

Capsule Long-term studies at two Scottish colonies show that the k value used to convert counts of individual Common Guillemots Uria aalge to pairs has changed substantially in the last 15 years due to decreases in survival and colony attendance. Any future wide-scale census of this species needs to collect concurrent k values if counts of individual birds are to be used to determine changes in breeding populations since the previous survey.

Intra-specific plasticity in parental investment in a long-lived single-prey loader

Seabirds exhibit considerable behavioural flexibility in foraging investment in order to meet the nutritional needs of their chicks during variable environmental conditions. Although regulation of offspring provisioning is generally thought to be related to species-specific constraints imposed by central place foraging, some studies suggest different responses within the same species linked to local differences in foraging conditions. Under adverse conditions, seabirds are expected to be less flexible because they must secure their own survival chances first before investing in current reproduction. Short-ranging single-prey loaders are expected to show large intra-specific variation in time spent on foraging because their mode of foraging is energetically expensive, and because they face restricted possibilities to increase the numerical prey input to the colony compared to multiple prey loaders. In this study, we examined if and how the single-prey loading Sandwich Tern Thalasseus sandvicensis varies colony attendance based on the nutritional status of their chick as well as parental body condition in two study colonies. The proportion of time that a chick was left unattended at the colony negatively correlated with chick body condition, suggesting that the parents tried to counterbalance poor feeding conditions by investing more time in foraging. Energy transport rates to the chicks (corrected for time spent away from the colony) and body condition of the chicks were similar in both colonies. However, at Zeebrugge, where adults were in poor body condition, parental non-attendance was much lower than on Griend, even when chicks were in poor condition. Still, our results suggest that parental nest non-attendance in Sandwich Terns is merely a corrective response to food loss to kleptoparasitic gulls in order to meet the nutritional status of the chick, although an effect of adult body condition could not be excluded.

Female Weddell seals show flexible strategies of colony attendance related to varying environmental conditions.

Many animal life cycles involve movements among different habitats to fulfill varying resource demands. There are inherent costs associated with such movements, and the decision to leave or stay at a given location ought to be motivated by the benefits associated with potential target habitats. Because movement patterns, especially those associated with reproduction, can have important implications for the success (survival, reproduction) of individual animals, and therefore a population's dynamics, it is important to identify and understand their sources of variation (environmental and individual). Here, using a mark-recapture, multistate modeling approach, we investigated a set of a priori hypotheses regarding sources and patterns of variation in breeding-colony attendance for Weddell seal (Leptonychotes weddellii) females on sabbatical from pup production. For such females, colony attendance might be motivated by predation avoidance and positive social interactions related to reproduction, but some costs, such as reduced foraging opportunities or aggressive interactions with conspecifics, might also exist. We expected these benefits and costs to vary with a female's condition and the environment. Results revealed that the probability of being absent from colonies was higher (1) in years when the extent of local sea ice was larger, (2) for the youngest and oldest individuals, and (3) for females with less reproductive experience. We also found substantial levels of residual individual heterogeneity in these rates. Based on our a priori predictions, we postulate that the decision to attend breeding colonies or not is directly influenced by an individual's physiological condition, as well as by the ice-covered distance to good foraging areas, availability of predator-free haul-out sites, and the level of negative interactions with conspecifics inside colonies. Our results support the idea that in iteroparous species, and colonial animals in particular, seasonal and temporary movements from/to reproductive sites represent flexible behavioral strategies that can play an important role in coping with environmental variability.

Carry-over body mass effect from winter to breeding in a resident seabird, the little penguin.

Using body mass and breeding data of individual penguins collected continuously over 7 years (2002–2008), we examined carry-over effects of winter body mass on timing of laying and breeding success in a resident seabird, the little penguin (Eudyptula minor). The austral winter month of July consistently had the lowest rate of colony attendance, which confirmed our expectation that penguins work hard to find resources at this time between breeding seasons. Contrary to our expectation, body mass in winter (July) was equal or higher than in the period before (‘moult-recovery’) and after (‘pre-breeding’) in 5 of 7 years for males and in all 7 years for females. We provided evidence of a carry-over effect of body mass from winter to breeding; females and males with higher body mass in winter were more likely to breed early and males with higher body mass in winter were likely to breed successfully. Sex differences might relate to sex-specific breeding tasks, where females may use their winter reserves to invest in egg-laying, whereas males use their winter reserves to sustain the longer fasts ashore during courtship. Our findings suggest that resident seabirds like little penguins can also benefit from a carry-over effect of winter body mass on subsequent breeding.

The Effect of Moonlight on Scopoli's Shearwater Calonectris diomedea Colony Attendance Patterns and Nocturnal Foraging: A Test of the Foraging Efficiency Hypothesis

AbstractMoonlight is known to affect the nocturnal behaviour and activity rhythms of many organisms. For instance, predators active at night may take advantage from increased visibility afforded by the moon, while prey might regulate their activity patterns to become less detectable. Many species of pelagic seabirds attend their colony only at night, in complete darkness, avoiding approaching their nest sites under moonlight. This behaviour has been most often interpreted as an antipredator adaptation (‘predation avoidance’ hypothesis). However, it may also reflect a lower foraging efficiency during moonlit nights (‘foraging efficiency’ hypothesis). Indeed, moonlight may reduce prey availability because preferred seabird prey is known to occur at higher depths in moonlit nights. Using high‐accuracy behavioural information from data loggers, we investigated the effect of moonlight on colony attendance and at‐sea nocturnal foraging in breeding Scopoli's shearwaters Calonectris diomedea. We found that birds departing for self‐feeding trips around the full moon performed longer trips than those departing around the new moon. On nights when the moon was present only partly, nest burrow entrances took place largely in the moonless portion of the night. Moreover, contrary to predictions from the ‘foraging efficiency’ hypothesis, nocturnal foraging activity increased according to moonlight intensity, suggesting that birds increased their foraging activity when prey became more detectable. This study strengthens the idea that colony attendance behaviour is strictly controlled by moonlight in shearwaters, which is possibly related to the perception of a predation risk.

Environmental correlates of temporary emigration for female Weddell seals and consequences for recruitment

In colonial‐breeding species, prebreeders often emigrate temporarily from natal reproductive colonies then subsequently return for one or more years before producing young. Variation in attendance–nonattendance patterns can have implications for subsequent recruitment. We used open robust‐design multistate models and 28 years of encounter data for prebreeding female Weddell seals (Leptonychotes weddellii [Lesson]) to evaluate hypotheses about (1) the relationships of temporary emigration (TE) probabilities to environmental and population size covariates and (2) motivations for attendance and consequences of nonattendance for subsequent probability of recruitment to the breeding population. TE probabilities were density dependent (βˆBPOP = 0.66, = 0.17; estimated effects [β] and standard errors of population size in the previous year) and increased when the fast‐ice edge was distant from the breeding colonies (βˆDIST = 0.75, = 0.04; estimated effects and standard errors of distance to the sea‐ice edge in the current year on TE probability in the current year) and were strongly age and state dependent. These results suggest that trade‐offs between potential benefits and costs of colony attendance vary annually and might influence motivation to attend colonies. Recruitment probabilities were greatest for seals that consistently attended colonies in two or more years (e.g., = 0.56, SD = 0.17) and lowest for seals that never or inconsistently attended prior to recruitment (e.g., = 0.32, SD = 0.15), where denotes the mean recruitment probability (over all years) for 10‐year‐old seals for the specified prebreeder state. In colonial‐breeding seabirds, repeated colony attendance increases subsequent probability of recruitment to the adult breeding population; our results suggest similar implications for a marine mammal and are consistent with the hypothesis that prebreeders were motivated to attend reproductive colonies to gain reproductive skills or perhaps to optimally synchronize estrus through close association with mature breeding females.

Movement models provide insights into variation in the foraging effort of central place foragers

Ecology and conservation depend on an understanding of how animals adjust their behaviour patterns in response to changes in their environment. Central place foragers (CPFs) are well-suited for developing ecological models of adaptive processes because their objective functions and operational constraints can be reasonably inferred. Central place foraging and provisioning theory provide the theoretical framework for this analysis. Analysis of CPF time allocation and energy budgets can provide insights into their strategies for responding to environmental variation. However, until recently, suitable high-resolution data on the behaviour of seabirds and other CPFs at sea have not been available. Previous studies of breeding seabirds have investigated variation in foraging trip duration and colony attendance, but few studies have analyzed variation in time allocation within foraging trips. Here, we develop a conceptual energy-based model for analysing variation in the time allocation of CPFs during foraging trips, and apply it to the movement patterns of Peruvian boobies (Sula variegata). Foraging trips of Peruvian boobies, recorded using high-resolution global positioning systems (GPS), were first partitioned into movement modes consistent with travel and foraging behaviours using a hidden Markov model (HMM) adapted to account for gaps in the GPS tracks associated with diving behaviour. Analysis of the HMM results based on the conceptual model indicated that differences in foraging effort between two treatments were best explained by a combination of differences in travel time and in time spent searching for prey. The conceptual model provides the basis for an integrated approach to analysis of variation in foraging strategies in which identification of various behaviours is coupled with assessments of time and energy budgets. This integrated approach can contribute to greater understanding of the processes determining foraging strategies and the limits to these strategies in the context of competition for resources and global climate change.

Colony attendance and at‐sea distribution of thin‐billed prions during the early breeding season

Procellariiform seabirds have extreme life histories; they are very long‐lived, first breed when relatively old, lay single egg clutches, both incubation and chick‐rearing are prolonged and chicks exhibit slow growth. The early part of the breeding season is crucial, when pair bonds are re‐established and partners coordinate their breeding duties, but is a difficult period to study in burrow‐nesting species. Miniature geolocators (Global Location Sensors or GLS loggers) now offer a way to collect data on burrow attendance, as well as determine at‐sea movements. We studied the early breeding season in thin‐billed prions Pachyptila belcheri breeding at New Island, Falkland Islands. Males and females arrived back at the colony at similar times, with peak arrival in the last days of September. However, males spent more time on land during the pre‐laying period, presumably defending and maintaining the burrow and maximising mating opportunities. Males departed later than females, and carried out a significantly shorter pre‐laying exodus. Males took on the first long incubation shift, whereas females returned to sea soon after egg laying. During the pre‐laying exodus and incubation, males and females travelled at similar speeds (> 250 km d−1) and were widely distributed over large areas of the Patagonian Shelf. Inter‐annual differences in oceanographic conditions were stronger during the incubation than during the pre‐laying exodus and were matched by stronger differences in distribution. The study thus suggests that extended trips and flexible distribution enable thin‐billed prions to meet the high energy demands of egg production and incubation despite low productivity in waters around the colony during the early summer.

Seasonal Sexual Segregation by Monomorphic Sooty Shearwaters Puffinus griseus Reflects Different Reproductive Roles during the Pre-Laying Period

Tracking technology has revolutionized knowledge of seabird movements; yet, few studies have examined sex differences in distribution and behavior of small to medium-sized, sexually-monomorphic seabirds. Application of bird-borne geolocation-immersion loggers revealed seasonal segregation in the sexually-monomorphic Sooty Shearwater Puffinus griseus, mainly in the pre-laying period, when there were clear differences in reproductive roles. Shearwaters first returned to the Falkland Islands on 27 Sept±8 d; males, on average, 8 d earlier than females. Prior to egg-laying, distribution at sea, colony attendance and behaviour depended on sex. Males foraged locally over the southern Patagonian Shelf and Burdwood Bank, spending mainly single days at sea and intervening nights in the burrow. Females, who flew for more of the day during this time, foraged in more distant areas of the northern Patagonian Shelf and Argentine Basin that were deeper, warmer and relatively more productive. Attendance of females at the colony was also more variable than that of males and, overall, males were present for significantly more of the pre-laying period (38 vs. 19% of time). Sex differences were reduced following egg-laying, with males and females using similar foraging areas and making trips of similar mean duration in incubation (7.6±2.7 d) and chick-rearing (1.4±1.3 d). Congruence continued into the non-breeding period, with both sexes showing similar patterns of activity and areas of occupancy in the NW Atlantic. Thus, seasonal changes in reproductive roles influenced patterns of sexual segregation; this occurred only early in the season, when male Sooty Shearwaters foraged locally, returning regularly to the colony to defend (or maintain) the burrow or the mate, while females concentrated on building resources for egg development in distant and relatively more productive waters.

A simulation model coupling the behaviour and energetics of a breeding central place forager to assess the impact of environmental changes

During the breeding season, seabirds are obligate central place foragers, and this may make them vulnerable to impacts of environmental change. An individual based model of a pair of central place foragers and their offspring has been developed for the common guillemot (Uria aalge). The behavioural decisions of each adult depend on the state of themselves, their partner and their chick. The behaviour of the adults and the body masses of all three are followed over the chick rearing period. The model was used to investigate how chick fledging mass, proportion of time the chick was left unattended by its parents and change in adult mass are impacted by different foraging ranges, prey abundance and calorific content. Adults in the model typically declined in mass during the chick rearing period, although none died of starvation. Provisioning parents can, to some extent, increase foraging distance from the colony without a reduction in the proportion of chicks reaching a suitable fledging mass or increasing the time they are left unattended. The foraging range at which a decline in fledging success and colony attendance occurs is influenced by changes in either one or both of, prey abundance and prey quality. Patterns produced by the model are consistent with field observations and biological knowledge. As the model outputs can give an indication of the fitness consequences of environmental changes it can be used to address theoretical ecological questions as well as to inform marine spatial management.