A goal of seabird ecology is to relate the physiology, population dynamics, distribution and behaviour of seabirds with their biotic and abiotic environments. One of the most challenging aspects is to understand how seabirds interact with their environment when direct observations are not always possible or practical. In the present paper, we reviewed 218 published studies that examined associations between seabird distribution, behaviour and their environment, in order to assess current trends, weaknesses and the future directions of research. Based on the number of publications, it is evident that the field is growing rapidly and that methods for evaluating seabird distribution are becoming increasingly more sophisticated and are changing from Eulerian (grid-like) to Lagrangian (particle-like) data types. This has been accompanied by a reduction in the spatial and temporal scale of observation, where, in most cases, no behavioural information is inferred from Lagrangian data; instead they are often used as if they were Eulerian data. In parallel, environmental remote sensing is becoming more common; however, we did not record significant changes in the statistical approaches used to describe seabird distributions and used to link them with oceanographic variables. In particular, despite the spatially explicit nature of the data, spatial statistics have rarely been used. The vast majority of studies used environmental variables that described water masses (descriptive approach), whereas a few studies determined oceanographic features that enhance prey availability to seabirds (process-based approach). Future studies could enhance their ecological interpretation of seabird-environment interactions by making greater use of ad hoc statistical approaches that facilitate appropriate pattern detection (e.g. area-restricted searching pattern for birds, mesoscale patterns for environment). Furthermore, appropriate hypothesis testing and modelling that accounts for the spatially explicit, multiscale and multivariate nature of the interaction between seabirds and their habitats is recommended. Although quantitative methods currently exist (but are rarely used), further application could greatly improve our understanding of the processes linking seabird distribution to their environment.

SUMMARY Interpolation of geolocation or Argos tracking data is a necessity for habitat use analyses of marine vertebrates. In a fluid marine environment, characterized by curvilinear structures, linearly interpolated track data are not realistic. Based on these two facts, we interpolated tracking data from albatrosses, penguins, boobies, sea lions, fur seals and elephant seals using six mathematical algorithms. Given their popularity in mathematical computing, we chose Bézier, hermite and cubic splines, in addition to a commonly used linear algorithm to interpolate data. Performance of interpolation methods was compared with different temporal resolutions representative of the less-precise geolocation and the more-precise Argos tracking techniques. Parameters from interpolated sub-sampled tracks were compared with those obtained from intact tracks. Average accuracy of the interpolated location was not affected by the interpolation method and was always within the precision of the tracking technique used. However, depending on the species tested, some curvilinear interpolation algorithms produced greater occurrences of more accurate locations, compared with the linear interpolation method. Total track lengths were consistently underestimated but were always more accurate using curvilinear interpolation than linear interpolation. Curvilinear algorithms are safe to use because accuracy, shape and length of the tracks are either not different or are slightly enhanced and because analyses always remain conservative. The choice of the curvilinear algorithm does not affect the resulting track dramatically so it should not preclude their use. We thus recommend using curvilinear interpolation techniques because of the more realistic fluid movements of animals. We also provide some guidelines for choosing an algorithm that is most likely to maximize track quality for different types of marine vertebrates.

Diving animals offer a unique opportunity to study the importance of physiological constraint and the limitation it can impose on animal's behaviour in nature. This paper examines the interaction between physiology and behaviour and its impact on the diving capability of five eared seal species (Family Otariidae; three sea lions and two fur seals). An important physiological component of diving marine mammals is the aerobic dive limit (ADL). The ADL of these five seal species was estimated from measurements of their total body oxygen stores, coupled with estimates of their metabolic rate while diving. The tendency of each species to exceed its calculated ADL was compared relative to its diving behaviour. Overall, our analyses reveal that seals which forage benthically (i.e. on the sea floor) have a greater tendency to approach or exceed their ADL compared to seals that forage epipelagically (i.e. near the sea surface). Furthermore, the marked differences in foraging behaviour and physiology appear to be coupled with a species demography. For example, benthic foraging species have smaller populations and lower growth rates compared to seal species that forage epipelagically. These patterns are relevant to the conservation and management of diving vertebrates.

Field metabolic rates (FMR) and activity patterns of black-browed albatrosses Thalassarche melanophrys were measured while at sea and on nest during the incubation stage at Kerguelen Island, southwestern Indian Ocean. Activity-specific metabolic rates of five albatrosses at sea (FMRat-sea) were measured using doubly labeled water (DLW), and by equipping birds with wet-dry activity data loggers that determined when birds were in flight or on the water. The metabolic rates of four birds incubating their eggs (FMRon-nest) were also measured using DLW. The mean+/-SD FMRat-sea of albatrosses was 611+/-96 kJ kg(-1) d(-1) compared to FMRon-nest of 196+/-52 0 kg(-1) d(-1). While at sea, albatrosses spent 52.9+/-8.2% (N = 3) of their time in flight and they landed on the water 41.2+/-13.9 times per day. The FMR of black-browed albatrosses appear to be intermediate to that of three other albatross species. Based on at-sea activity, the power requirement of flight was estimated to be 8.7 W kg(-1) (or 4.0 x predicted BMR), which is high compared to other albatross species, but may be explained by the high activity levels of the birds when at sea. The FMRat-sea of albatrosses, when scaled with body mass, are lower than other seabirds of similar body size, which probably reflects the economical nature of their soaring flight.

Wandering AlbatrossDiomedea exulanschicks require 9–10 months to achieve adult body size at fledging, at which time they are also sexually size dimorphic. Because the developmental period spans the winter season, chicks must endure severe winter conditions and variability in provisioning effort by their parents. Thus chicks may adjust their rate of energy utilization to accommodate variations in provisioning, but this has not previously been studied. We followed longitudinally the changes in growth, body composition and oxygen consumption of 10 chicks from the end of the brooding period until fledging on the Crozet Islands. Body mass, culmen length and wing length were measured every 10 days and total body water (TBW) and resting metabolic rate (RMR) were measured monthly. Overall growth followed a logistic curve for all chicks, and sexual dimorphism in body mass appeared as early as the second month of measurements (males being heavier than females). Absolute TBW followed a logistic increase like that of body mass and was significantly higher in males owing to the difference in body mass. Conversely, mass‐specific TBW (i.e. the proportion of body mass made up of water) did not differ significantly between male and female chicks. Absolute RMR peaked at 1.5 × adult basal metabolism in midwinter when chicks achieved maximum body mass, but decreased to adult levels by the time chicks fledged. The decrease in absolute RMR following attainment of peak mass is atypical of most seabird chicks (Procellariiformes) and may be explained partly by a reduction in size of the gut when parents reduce provisioning effort. The changes in mass‐specific RMR did not differ between sexes but male chicks, being heavier, had higher absolute oxygen consumption and therefore greater energy requirements.

SUMMARY The period of maternal dependence is a time during which mammalian infants must optimise both their growth and the development of behavioural skills in order to successfully meet the demands of independent living. The rate and duration of maternal provisioning, post-weaning food availability and climatic conditions are all factors likely to influence the growth strategies of infants. While numerous studies have documented differences in growth strategies at high taxonomic levels, few have investigated those of closely related species inhabiting similar environments. The present study examined the body composition, metabolism and indices of physiological development in pups of Antarctic fur seals (Arctocephalus gazella) and subantarctic fur seals (Arctocephalus tropicalis), congeneric species with different weaning ages (4 months and 10 months, respectively), during their overlap in lactation at a sympatric breeding site in the Iles Crozet. Body lipid reserves in pre-moult pups were significantly greater (t28=2.73, P<0.01) in subantarctic (26%) than Antarctic fur seals (22%). Antarctic fur seal pups, however, had significantly higher (t26=3.82, P<0.001) in-air resting metabolic rates (RMR; 17.1±0.6 ml O2 kg-1 min-1) than subantarctic fur seal pups (14.1±0.5 ml O2 kg-1 min-1). While in-water standard metabolic rate (SMR; 22.9±2.5 ml O2 kg-1 min-1) was greater than in-air RMR for Antarctic fur seal pups (t9=2.59, P<0.03), there were no significant differences between in-air RMR and in-water SMR for subantarctic fur seal pups (t12=0.82, P>0.4), although this is unlikely to reflect a greater ability for pre-moult pups of the latter species to thermoregulate in water. Pup daily energy expenditure was also significantly greater (t27=2.36, P<0.03) in Antarctic fur seals (638±33 kJ kg-1 day-1) than in subantarctic fur seals (533±33 kJ kg-1 day-1), which corroborates observations that pups of the former species spend considerably more time actively learning to swim and dive. Consistent with this observation is the finding that blood oxygen storage capacity was significantly greater (t9=2.81, P<0.03) in Antarctic (11.5%) than subantarctic fur seal (8.9%) pups. These results suggest that, compared with subantarctic fur seals, Antarctic fur seal pups adopt a strategy of faster lean growth and physiological development, coupled with greater amounts of metabolically expensive behavioural activity, in order to acquire the necessary foraging skills in time for their younger weaning age.

Summary Theoretical models predict that animals will vary their effort to maximize different currencies such as time and energy when the constraints of reproduction change during breeding, but this has been poorly studied in free‐ranging animals. Foraging effort (energy per unit time) was examined by comparing mass changes, foraging costs and activity‐specific behaviours of Wandering Albatrosses (Diomedea exulans Linnaeus) during the incubation and chick‐brooding stages. In 1998, 38 albatrosses (20 during incubation and 18 during brooding) were injected with doubly labelled water and equipped with satellite transmitters and activity data loggers. During incubation, albatrosses travelled 3·7 times farther and were at sea 3·2 times longer, yet foraging costs were significantly lower than trips made during brooding (incubation 4·52 ± 0·50 SD W kg−1vs brooding 4·98 ± 0·55 SD W kg−1). The rate of daily mass gain decreased significantly with time at sea during incubation whereas the rate of daily mass gain increased significantly with time at sea during brooding. Foraging effort was higher during brooding, suggesting that birds were minimizing time at sea to maximize the rate of food delivery to chicks. In contrast, foraging effort was lower during incubation, suggesting that birds were maximizing time at sea and minimizing the energy costs of foraging. Foraging costs were also different between sexes. However, this was related to body size differences and not to differences in foraging effort as suggested in previous studies.

We studied the changes in heart rate (HR) associated with metabolic rate of incubating and resting adult wandering albatrosses (Diomedea exulans) on the Crozet Islands. Metabolic rates of resting albatrosses fitted with external HR recorders were measured in a metabolic chamber to calibrate the relationship between HR and oxygen consumption (V(O(2))) (V(O(2))=0.074 x HR+0.019, r(2)=0.567, P<0.001, where V(O(2)) is in ml kg(-1) min(-1) and HR is in beats min(-1)). Incubating albatrosses were then fitted with HR recorders to estimate energy expenditure of albatrosses within natural field conditions. We also examined the natural variation in HR and the effects of human disturbance on nesting birds by monitoring the changes in HR. Basal HR was positively related to the mass of the individual. The HR of incubating birds corresponded to a metabolic rate that was 1.5-fold (males) and 1.8-fold (females) lower than basal metabolic rate (BMR) measured in this and a previous study. The difference was probably attributable to birds being stressed while they were held in the metabolic chamber or wearing a mask. Thus, previous measurements of metabolic rate under basal conditions or for incubating wandering albatrosses are likely to be overestimates. Combining the relationship between HR and metabolic rate for both sexes, we estimate that wandering albatrosses expend 147 kJ kg(-1) day(-1) to incubate their eggs. In addition, the cost of incubation was assumed to vary because (i) HR was higher during the day than at night, and (ii) there was an effect of wind chill (<0 degrees C) on basal HR. The presence of humans in the vicinity of the nest or after a band control was shown to increase HR for extended periods (2-3 h), suggesting that energy expenditure was increased as a result of the disturbance. Lastly, males and females reacted differently to handling in terms of HR response: males reacted more strongly than females before handling, whereas females took longer to recover after being handled.

Summary The study examined the behavioural factors that influence variation in foraging costs of wandering albatrosses (Diomedea exulans Linnaeus) during two consecutive years. This was performed by measuring simultaneously foraging activities and metabolic rates (W kg−1) of breeding birds during the incubation periods of 1998 and 1999. A total of 38 birds (19 in each year) were injected with doubly labelled water, equipped with satellite transmitters to measure foraging distance and flight speed, and equipped with activity recorders to measure landings and take‐offs from the water surface. In 1999, birds spent on average 30% more time at sea (11·07 ± 3·53 SD, days), travelled 43% further (5473 ± 2348 SD, km) and increased the number of landings and take‐offs per day by 66% (28·1 ± 11·4 SD, landings per day) compared to 1998. Energy expenditure rates during foraging were significantly higher in 1999 (5·3 ± 0·8 SD, W kg−1, n = 8) compared to 1998 (4·5 ± 0·5 SD, W kg−1, n = 11). Landing and take‐off rates explained the greatest proportion of variation in energy expenditure rates of wandering albatrosses, rather than total distance travelled, flight speed or time in flight. Despite 18% higher foraging costs in 1999, the daily rate of food intake was similar between years, suggesting that energy acquisition per unit effort was lower. The energetic cost of flight for wandering albatrosses was estimated to fall between 3·4 and 5·1 W kg−1, or 1·4–2·0 × measured BMR, one of the lowest flight cost yet reported for any seabird.

Measurements of incubation energetics can vary depending on the method used to measure metabolism of an incubating bird. Therefore, we evaluated the energy expenditure of six male and four female wandering albatrosses (Diomedea exulans Linnaeus) using doubly labeled water (DLW), the rate of mass loss, and estimates of metabolic water production derived from water influx rate (WIR). Incubation metabolic rates (IMR) determined with DLW (169 +/- 21 kJ kg(-1) d(-1) SD) were significantly lower than estimates derived from mass loss (277 +/- 46 kJ kg(-1) d(-1) SD) and WIR (males = 289 +/- 60 kJ kg(-1) d(-1) vs. females = 400 +/- 69 kJ kg(-1) d(-1) SD). Estimates of IMR from mass loss and WIR were similar to IMR (305 +/- 39 kJ kg(-1) d(-1) SD) determined by respirometry in a previous study, and IMR from DLW was similar to estimates based on heart rate(HR;147 +/- 26 kJ kg(-1) d(-1) SD) determined in another study. Applying the different measurements of IMR to construct an energy budget, we estimate that a breeding pair of wandering albatrosses spends 124-234 MJ to incubate the egg for 78 d. Finally, IMRs determined with DLW and HR were similar to estimated basal metabolic rates derived from six different allometric equations, suggesting that heat production from adult maintenance metabolism is sufficient to incubate the egg.

1. The investigation covered whether sexual dimorphism could affect flight performance in a manner that is consistent with differences in at-sea distribution of male and female Wandering Albatrosses, Diomedea exulans Linnaeus. Adult morphology was also compared to near-fledged chicks to assess whether morphological differences are consistent with different at-sea distributions of adults and fledglings.2. Body girth, mass, wing span and area were measured on 24 females (16 adults and 8 chicks) and 32 males (20 adults and 12 chicks) breeding in the Crozet Archipelago.3. On average, adult males had longer wings (4%, 311 +/- 4 cm) with 6.8% more area (6260 +/- 270 cm(2)), but were also 20.4% heavier (9.44 +/- 0.59 kg) than adult females. As a result, wing loading in adult males was 12.1% greater than adult females.4. When compared with adults, total wing area of chicks was lower resulting in higher wing loading because of the incomplete growth of chicks at the time measurements were collected. However, projected chick growth to fledging indicates that wing loading would be lower in fledglings than adults.5. Because wing loading determines flight speed, it is conceivable that windier regions of the sub-Antarctic/Antarctic are more optimal for male albatrosses because they have higher wing loading. Conversely, wing loading is lower in adult females and fledglings, which could make them better adapted to exploit lighter winds of the subtropical and tropical regions. Thus, size dimorphism may have a functional role in flight performance that influences the at-sea distribution of adult and fledgling Wandering Albatrosses.

The influence of wind patterns on behaviour and effort of free-ranging male wandering albatrosses (Diomedea exulans) was studied with miniaturized external heart-rate recorders in conjunction with satellite transmitters and activity recorders. Heart rate was used as an instantaneous index of energy expenditure. When cruising with favourable tail or side winds, wandering albatrosses can achieve high flight speeds while expending little more energy than birds resting on land. In contrast, heart rate increases concomitantly with increasing head winds, and flight speeds decrease. Our results show that effort is greatest when albatrosses take off from or land on the water. On a larger scale, we show that in order for birds to have the highest probability of experiencing favourable winds, wandering albatrosses use predictable weather systems to engage in a stereotypical flight pattern of large looping tracks. When heading north, albatrosses fly in anticlockwise loops, and to the south, movements are in a clockwise direction. Thus, the capacity to integrate instantaneous eco-physiological measures with records of large-scale flight and wind patterns allows us to understand better the complex interplay between the evolution of morphological, physiological and behavioural adaptations of albatrosses in the windiest place on earth.

The white whale Delphinapterus leucas is an exceptional diver, yet we know little about the physiology that enables this species to make prolonged dives. We studied trained white whales with the specific goal of assessing their diving and swimming performance. Two adult whales performed dives to a test platform suspended at depths of 5-300 m. Behavior was monitored for 457 dives with durations of 2.2-13.3 min. Descent rates were generally less than 2 m s(-1) and ascent rates averaged 2.2-3 m s(-1). Post-dive plasma lactate concentration increased to as much as 3.4 mmol l(-1) (4-5 times the resting level) after dives of 11 min. Mixed venous PO2 measured during voluntary breath-holds decreased from 79 to 20 mmHg within 10 min; however, maximum breath-hold duration was 17 min. Swimming performance was examined by training the whales to follow a boat at speeds of 1.4-4.2 m s(-1). Respiratory rates ranged from 1.6 breaths min(-1) at rest to 5.5 breaths min(-1) during exercise and decreased with increasing swim speed. Postexercise plasma lactate level increased to 1.8 mmol l(-1) (2-3 times the resting level) following 10 min exercise sessions at swimming speeds of 2.5-2.8 m s(-1). The results of this study are consistent with the calculated aerobic dive limit (O-2 store/metabolic rate) of 9-10 min. In addition, white whales are not well adapted for high-speed swimming compared with other small cetaceans.