Adult Brünnich's Guillemots Uria lomvia balance body condition and investment in chick growth

To investigate the covariation of adult body condition and nestling growth, we weighed adult Brünnich's Guillemots Uria lomvia rearing chicks at Coats Island, Nunavut, Canada, each year between 1988 and 2002. We estimated chick mass at 14 days for a sample of chicks reared in the same years. Adult mass and chick mass at 14 days were highly correlated, suggesting that, as feeding conditions deteriorate, adults compromise by reducing their own body reserves, while at the same time delivering less food to their offspring. We compared the prediction of the least-squares regression for the Coats Island data with observations made at Digges Island, a much larger colony about 300 km away, where birds are similar in linear body measurements to those at Coats Island and have a similar body mass while incubating. Adult mass at Digges Island averaged 11% less during chick-rearing than during incubation, compared with only a 5% difference at Coats Island. Mean chick mass at 14 days at Digges Island was lower in all years than was observed for chicks at Coats Island in any year. The observed 14-day chick masses at Digges Island in two years were close to values predicted by adult mass and somewhat lower in two other years (those when chick growth was slowest). At Digges Island, the distribution of mass for brooding adults was right skewed and suggested a lower threshold at 800–850 g, below which Brünnich's Guillemots terminate breeding. We conclude that the correlation between adult and chick mass represents a dynamic equilibrium in which adults simultaneously adjust their own energy reserves and their delivery rate to the chick. This compromise must be based on behavioural choices made by individual birds and is unlikely to be a passive consequence of fluctuating conditions.     

Social interactions among breeding Brünnich's Guillemots Uria lomvia suggest constraints in relation to offspring vulnerability

Brünnich's Guillemots Uria lomvia are adapted to high-density breeding in large colonies on steep cliffs. Because they breed on narrow ledges, egg loss through dislodgement is an important cause of breeding failure. Fighting among breeders presumably raises the risk of accidental dislodgement. In this study, we investigated whether social behaviour among Brünnich's Guillemots shows any adaptations to reduce accidental egg loss by modifying behaviour during incubation. We found that the amount of aggression increased significantly at the time of hatching, perhaps in response to the reduced risk of breeding failure through dislodgement of a chick, compared with an egg. Allopreening followed an inverse trend, falling significantly after the day of hatch. This supports the hypothesis that allopreening is used to reduce aggressive interactions. At the same time, the frequency of allopreening was greatly increased on days when mosquitoes affected the birds, consistent with the hypothesis that allopreening is part of a defence strategy against ectoparasites.     

Sex-biased aggression and male-only care at sea in Brünnich's Guillemots Uria lomvia and Razorbills Alca torda

In several groups in the order Charadriformes, biparental care is followed by a period of male‐only care. Several hypotheses attempting to explain extended male parental care in shorebirds do not fit the Alcini. In a previous study of Brünnich’s Guillemots Uria lomvia and Razorbills Alca torda, we did not find support for female‐biased parental effort at the breeding site that would lead to males being in better condition to care for chicks at sea. However, in both species, males spent more off‐duty time at the breeding site than females, suggesting greater involvement in the defence of egg or chick, breeding site and mate. We predicted that there would be a male bias in size and aggressive behaviour associated with parental roles. To test this, body size and aggression of attending male and female Brünnich’s Guillemots and Razorbills were measured during incubation and brooding on the Gannet Islands, Labrador. Parental aggression was measured using natural observations of all agonistic interactions and, in Razorbills only, in situ responses to presentations of a predator model. In both species, males were significantly larger than females in culmen and gape length. Guillemot males initiated agonistic interactions more frequently than females during incubation. In contrast, female Guillemots were subjected to aggression more frequently than males and as a result were involved in more fights. In addition, the few chicks that were seen to die were being attended by single females. During the brooding period, Razorbill males responded aggressively to intruders more frequently than females, made more aggressive responses than females, and responded aggressively more frequently and more intensely than females to a predator model. In both species there was a similar male bias in morphology and behaviour that is consistent with male parents being more capable of protecting their chick, a probable advantage to chick survival during the uniparental care phase of some Charadriformes.     

Body mass changes in Brünnich's guillemots Uria lomvia with age and breeding stage

Body mass of Brünnich's guillemots Uria lomvia breeding at Coats Island, Canada, was measured during incubation and chick-rearing in 1988–2001. In most years, mass increased during incubation and fell after hatching, leveling off by the time chicks were 18 d old, close to the age at which chicks departed. Mass during incubation increased with age up to about 12 yr, but the mass of birds brooding chicks was not related to age. The trend towards increasing mass during incubation was mainly a consequence of mass increases of young breeders as older birds maintained a constant mass. The variation in adult mass with age during incubation seems likely to reflect age-related variation in foraging ability, but the loss of mass after hatching, being greater for older birds, appears best explained as a response to the demands of provisioning chicks, with older birds transferring their accumulated reserves to their chicks via higher provisioning rates.     

Seasonal declines in incubation periods of Brünnich's Guillemots Una lomvia:testing proximate causes

In many species of bird, eggs laid late in the laying period hatch after a shorter incubation period than do eggs laid early. These seasonal declines in incubation period are generally thought to confer evolutionary advantages, but the proximate mechanisms that underlie them are poorly understood. Seasonal declines in incubation period are usually attributed to: (1) seasonal increases in ambient air temperatures; (2) seasonal changes in the behaviour of incubating birds; and/or (3) seasonal declines in egg size. In a previous study, Common Guillemot Una aalge incubation periods declined with laying date at a low-Arctic colony. As there was no support for hypotheses 1 or 2, it was suggested that this occurred because egg size declined with laying date, but eggs were not measured in that study. We recorded similar seasonal declines in the incubation periods of the single eggs laid by Brünnich's Guillemots Una lomvia at two low-Arctic colonies in four years. Neither seasonal variation in ambient air temperatures, nor in the behaviour of incubating adults, appeared to cause the declines. As predicted for Common Guillemots, incubation period increased with egg size among Briinnich's Guillemots, in one of two years. However, incubation period declined with laying date in the absence of corresponding declines in egg size. We conclude that none of the three commonly proposed proximate mechanisms adequately explains the seasonal variation in guillemot incubation periods. Several testable, alternative mechanisms are explored.     

Water temperature sampling by foraging Brünnich's Guillemots with bird-borne data loggers

We describe the features of waters where seabirds were feeding by sampling vertical water temperature profiles with data loggers mounted on five Brünnich's Guillemots in Svalbard, Norway. The guillemots foraged in a cold water (−0.5–0.5°C SST (sea surface temperature)) by making 1.8 dive bouts in short trips (32–257 min duration) as well as in moderate (0.5–2.0°C SST) and warm waters (2.5–4.0°C SST) by making 6.0 dive bouts during long trips (411–688 min duration). Judging from outbound flying time (15.7–24.4 min), time between dive bouts (23.9–43.3 min) and water types, the birds probably fed in fjord or coastal waters during short trips and in both coastal and offshore waters during long trips. Water temperature and diving behaviour can be simultaneously recorded by small data loggers, which therefore will provide useful information on marine features and foraging activity of top predators.     

Fatty acid signatures of female Brünnich's guillemots Uria lomvia suggests reliance on local prey for replacement egg production

The laying of smaller replacement eggs has been described as a time-saving adaptation because chicks generally grow faster once hatched than inside the egg. However, chicks hatched from smaller eggs have, potentially, lower survival. Consequently, the smaller replacement egg represents a benefit to the female in terms of preserving its own condition at a cost to their offspring. We test these ideas by measuring adult mass changes and plasma lipid concentration changes in male and female Brünnich's guillemots Uria lomvia breeding on Coats Island, Nunavut. Though males lost more mass than females, these differences were not significant. Between laying the first and replacement egg, plasma fatty acid concentrations declined in females and increased in males, suggesting that females mobilise less lipid to preserve their condition after laying the replacement egg. In females, plasma lipid concentrations of the dominant fatty acids found in the eggs (16:0 and 18:1) declined between the laying of first and replacement eggs while plasma concentrations of 20:1 increased in both males and females. We compared the fatty acid signatures of first and replacement egg to look for evidence of differences between the lipid sources for their production. Principal component and discriminant function analyses showed that the fatty acid signatures of replacement eggs were closer to the signatures of the local prey than those of first eggs. We suggest that females rely on local sources of energy to a greater degree for the production of the replacement egg than the first egg, but that endogenous reserves of certain nutrients are important for the production of both eggs.     

Body mass changes in Brünnich's guillemots Uria lomvia with age and breeding stage

Body mass of Brünnich's guillemots Uria lomvia breeding at Coats Island, Canada, was measured during incubation and chick‐rearing in 1988–2001. In most years, mass increased during incubation and fell after hatching, leveling off by the time chicks were 18 d old, close to the age at which chicks departed. Mass during incubation increased with age up to about 12 yr, but the mass of birds brooding chicks was not related to age. The trend towards increasing mass during incubation was mainly a consequence of mass increases of young breeders as older birds maintained a constant mass. The variation in adult mass with age during incubation seems likely to reflect age‐related variation in foraging ability, but the loss of mass after hatching, being greater for older birds, appears best explained as a response to the demands of provisioning chicks, with older birds transferring their accumulated reserves to their chicks via higher provisioning rates.     

Effects of prey abundance on the foraging behaviour, diving efficiency and time allocation of breeding Guillemots Uria aalge

The foraging behaviour of Guillemots Uria aalge at sea was compared between 2 years of radically different food abundance. Radio telemetry was used to determine foraging locations and diving patterns. In the poor compared with the good food year, foraging trips were much longer, the birds foraged more than six times further from their breeding sites, they spent over five times as much time diving when at sea and their estimated energy expenditure was twice as great. Time spent foraging in the poor food year was at the expense of time spent sitting at the colony. The duration of a foraging trip was a poor indicator of distance travelled but a good indicator of the amount of time spent diving. Mean dive durations, surface pause durations and interbout periods did not differ between years, but individuals made more than four times as many dives per diving bout in the poor food year. Surface pause lengths did not vary with water depth in either year. In the poor food year, birds made shorter surface pauses for a dive of a gi^ven duration than in the good food year, possibly accepting a lactic acid debt in order to maximize searching time, The duration of the interbout period was positively related to the number of dives in the previous bout, and dives tended to get shorter in long diving sequences, suggesting possible exhaustion effects. These data demonstrate that breeding Guillemots have the capacity to adjust their foraging behaviour and time budgets in response to changes in food abundance, but this flexibility was not sufficient to compensate fully for the very low food abundance experienced by birds in this study.     

Intercolony movement and behaviour of immature Guillemots Uria aalge

A minimum of 61 Guillemots Uria aalge ringed as chicks at other colonies was recorded at the Isle of May between 1987 and 1991 (four prior to 1990, six in 1990, 51 in 1991). Two were adults and 59 were immatures. Most of the British and Irish colonies where many chicks had been ringed were represented. Higher proportions of birds ringed at colonies relatively close to the Isle of May were observed compared to birds ringed at more distant sites. Visitors ranged in age from 2 to 5 years, with single 10- and 14-year-olds. Relatively fewer 2- and 5- than 3- and 4-year-old birds were seen. Visiting birds were usually recorded once only, significantly less often than native immatures of the same age, except for 2-year-olds. The proportion of birds occupying sites on intertidal rocks as against sites in the colony appeared to be higher compared with native birds of the same age. Visiting birds formed a substantial proportion of the immature population. Ten immature birds from the Isle of May were recorded elsewhere. Four of these had been seen on the Isle of May previously, and four were seen there subsequently. A 6-year-old hatched at Sumburgh ( c. 420 km by sea) bred at the Isle of May in 1992.     

An application of optimal diving models to diving behaviour of Brünnich's guillemots

Although theoretical models predict that the quality of foraging patches has little effect on optimal dive time with increasing depth, many empirical studies show that dive time at a given depth may vary. We developed a model that incorporated patch quality as a parameter of energy intake as a nonlinear function of time, and applied it to the diving behaviour of Brünnich's guillemots, Uria lomvia. The model indicated that optimal dive time can vary widely depending on the parameter. It also explained the convergence of observed dive times with travel time. Assuming the birds dived optimally, this parameter can be estimated from travel time and dive time for each dive. Foraging patches with larger estimated parameter values were favoured by the birds, suggesting that the parameter indicated patch quality. We used this parameter to test an optimal patch use model in divers. The results indicate that Brünnich's guillemots adjust their diving behaviour adaptively depending on patch quality, and that the optimal diving model is valid for prediction of observed dive patterns if patch quality is incorporated appropriately. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

The visual fields of Common Guillemots Uria aalge and Atlantic Puffins Fratercula arctica: foraging,vigilance and collision vulnerability

Significant differences in avian visual fields are found between closely related species that differ in their foraging technique. We report marked differences in the visual fields of two auk species. In air, Common Guillemots Uria aalge have relatively narrow binocular fields typical of those found in non‐passerine predatory birds. Atlantic Puffins Fratercula arctica have much broader binocular fields similar to those that have hitherto been recorded in passerines and in a penguin. In water, visual fields narrow considerably and binocularity in the direction of the bill is probably abolished in both auk species. Although perceptual challenges associated with foraging are similar in both species during the breeding season, when they are piscivorous, Puffins (but not Guillemots) face more exacting perceptual challenges when foraging at other times, when they take a high proportion of small invertebrate prey. Capturing this prey probably requires more accurate, visually guided bill placement and we argue that this is met by the Puffin's broader binocular field, which is retained upon immersion; its upward orientation may enable prey to be seen in silhouette. These visual field configurations have potentially important consequences that render these birds vulnerable to collision with human artefacts underwater, but not in air. They also have consequences for vigilance behaviour.     

Within- and between-year variation in the juvenile survival of Common Guillemots Uria aalge

We studied juvenile survival of 20 cohorts of Common Guillemot Uria aalge chicks colour-ringed on the Isle of May, Scotland, using both live observations at the colony and dead recoveries, allowing estimation of fidelity to the colony as well as survival. In this seabird, chicks leave the colony when only partly grown and are cared for by the male parent for several weeks afterwards. First-year survival varied strongly between cohorts, with a mean of 56% (range 30–91%). We did not identify any covariates which could explain this variation, whether relating to climate, population size or prey density. Survival was low during two regime shift episodes in the North Sea (1987–90 and 2000 onwards). Early hatched chicks were substantially more likely to survive than those hatching later in most years, whereas body condition at ringing had no detectable effect. Ringing recoveries indicated that mortality was highest in mid-winter, i.e. well after the cessation of paternal care. These results do not support the hypothesis that variation in prey quantity or energy content before fledging is a primary driver of variation in juvenile survival. Rather, it seems that chicks of high-quality parents are more likely to survive, as high-quality females tend to lay earlier in the season, and high-quality males presumably are better able to prepare their chicks to survive their first winter at sea. Very few (4%) Guillemots emigrated permanently before age 3 years, but from age 5 onwards 25–30% of birds annually left the colony or otherwise became unobservable.     

Foraging behaviour and time allocation of chick-rearing Razorbills Alca torda at Græsholmen, central Baltic Sea

A new type of bird-borne data logger, which stores data from flight and depth sensors at pre-set intervals, was used to investigate the foraging pattern and diving behaviour of chick-rearing Razorbills Alca torda breeding on the islet of Græsholmen (central Baltic Sea, Denmark). The instruments recorded all relevant events in the 35 foraging trips accomplished by six different individual birds; a seventh bird, equipped with a direction recorder, provided information on directional preferences exhibited on seven different trips. A foraging trip usually consisted of a number of flights interrupted by a small number of dives probably performed to explore the site for prey availability. True foraging occurred during the last stops in the outbound trip, after which the bird returned to the nest with a single flight or a sequence of a few flights. The duration of nocturnal trips was significantly longer than diurnal trips, possibly due to a preference for the familiar marine environment shown by birds when they are not 'on duty' in the nest. The majority of dives consisted of V-shaped dives in which birds descend to a maximum depth and then start ascending. A clear diurnal variation of the dive depth, which never exceeded 43 m, was recorded. Our results show that the new types of recording devices can be used to answer basic questions about the foraging strategies of marine birds.     

Among‐colony synchrony in the survival of Common Guillemots Uria aalge reflects shared wintering areas

Spatiotemporal variation in survival may be an important driver of multi‐population dynamics in many wild animal species, yet few scientific studies have addressed this issue, primarily due to a lack of sufficiently comprehensive and detailed datasets. Synchrony in survival rates among different, often distant, subpopulations appears to be common, caused by spatially correlated environmental conditions or by movement of animals from different sites such that their ranges overlap. Many seabird populations are effectively isolated during the breeding season because colonies are widely separated, but over the winter, birds disperse widely and there may be much mixing between different populations. The non‐breeding season is also the period of main mortality for seabirds. Using mark–recapture and ring‐recovery data, we tested for spatial, temporal and age‐related correlations in survival of Common Guillemots Uria aalge among three widely separated Scottish colonies that have varying overlap in their overwintering distributions. Survival was highly correlated over time for colonies/age‐classes sharing wintering areas and, except in 2004, was essentially uncorrelated for those with separate wintering areas. These results strongly suggest that one or more aspects of the winter environment are responsible for spatiotemporal variation in survival of British Guillemots, and provide insight into the factors driving large‐scale population dynamics of the species.     

Consequences of foraging trip duration on provisioning behaviour and fledging condition of common murres Uria aalgae

We examine the provisioning constraints of a pursuit‐diving seabird in a cold ocean regime by comparing the behaviour of common murres Uria aalge rearing chicks at two colonies in the Northwest Atlantic during 1998‐2000. Funk Island is the largest (340,000–400,000 breeding pairs) and most offshore (60 km) colony of common murres in eastern Canada. Seventy‐five percent of the Northwest Atlantic population of common murres breeds on this island. Great Island is one island within the Witless Bay Ecological Reserve, which is the second largest breeding aggregation (100 000 breeding pairs) and is located near‐shore (2 km). The primary forage fish species in Newfoundland waters is capelin Mallotus villosus, which spawns on or near coastal beaches during summer. Therefore, the two study colonies differ in their distance to food resources and colony size. It is within this natural context that we compare: (1) prey types and frequency of delivery (amount of prey), (2) parental time budgets, and (3) the mass and condition (mass/wing length) of fledglings at both colonies. Similarly sized female capelin (100–150 mm) were delivered to chicks at both colonies. Foraging time per day per parent, a proxy of foraging effort, was similar at both colonies (Great Island: 5.1 h; Funk Island: 5.5 h), as was the percentage of time spent with mates (Great Island: 12.3%; Funk Island: 10.9%). Foraging trips, however, were longer at Funk Island (4.1 h) than at Great Island (2.9 h). This resulted in lower feeding rates of chicks (0.17 feeds per h) and poorer condition of fledglings (2.9 g/mm) at Funk Island compared to those at Great Island (0.22 feeds per h; 3.9 g/mm). We hypothesize that provisioning efforts are constrained at Funk Island by (1) distant food resources and increased competitor density, resulting in longer foraging trip durations and (2) the time spent paired with mates at the colony, which may reflect a minimum time required to maintain breeding sites due to higher breeding densities at Funk Island compared to Great Island. Demographic consequences of this poor fledgling condition at Funk Island are unknown, but fledglings may sufficiently accelerate growth at sea due to their closer proximity to an important nursery area. If fledgling survival is compromised, however, the lower potential for growth at Funk Island will impact the entire Northwest Atlantic population of common murres.