Determining energy expenditure in a large seabird using accelerometry.

The trade off between energy gained and expended is the foundation of understanding how, why and when animals perform any activity. Based on the concept that animal movements have an energetic cost, accelerometry is increasingly being used to estimate energy expenditure. However, validation of accelerometry as an accurate proxy for field metabolic rate in free-ranging species is limited. In the present study, Australasian gannets (Morus serrator) from the Pope's Eye colony (38°16'42″S 144°41'48″E), south-eastern Australia, were equipped with GPS and tri-axial accelerometers and dosed with doubly labelled water (DLW) to measure energy expenditure during normal behaviour for 3-5 days. The correlation between daily energy expenditure from the DLW and vectorial dynamic body acceleration (VeDBA) was high for both a simple correlation and activity-specific approaches (R2=0.75 and 0.80, respectively). Varying degrees of success were observed for estimating at-sea metabolic rate from accelerometry when removing time on land using published energy expenditure constants (R2=0.02) or activity-specific approaches (R2=0.42). The predictive capacity of energy expenditure models for total and at-sea periods was improved by the addition of total distance travelled and proportion of the sampling period spent at sea during the night, respectively (R2=0.61-0.82). These results indicate that accelerometry can be used to estimate daily energy expenditure in free-ranging gannets and its accuracy may depend on the inclusion of movement parameters not detected by accelerometry.

Temporal and spatial isotopic variability of marine prey species in south-eastern Australia: Potential implications for predator diet studies.

Stable isotope analyses, particularly of carbon (δ13C) and nitrogen (δ15N), are used to investigate ecological relationships among species. For marine predators, research has shown the main factors influencing their intra-specific and intra-individual isotopic variation are geographical movements and changes in the composition of diet over time. However, as the differences seen may be the result of changes in the prey items consumed, a change in feeding location or the combination of both, knowledge of the temporal and spatial consistency in the isotopic values of prey becomes crucial for making accurate inferences about predator diets. This study used an abundant marine predator, the Australasian gannet (Morus serrator), as prey sampler to investigate the annual variation in fish and squid prey isotope values over a four-year period (2012-2015) and the geographic variation between two sites with contrasting oceanographic conditions. Significant inter-annual variation was observed in δ13C and/or δ15N values of five of the eight prey species analysed. The strongest inter-annual variation in both δ13C and δ15N values occurred in 2015, which coincided with a strong El Niño-Southern Oscillation (ENSO). This may suggest a temporal fluctuation in the geographic source of prey or the origin of their nutrients. These results suggest that it is important to consider the potential significant differences in isotopic values within the prey assemblages that predators consume. This is important to improve the interpretation of marine predator isotope results when determining the influence of environmental variability on their diets.

Geographical, temporal and individual factors influencing foraging behaviour and consistency in Australasian gannets.

Foraging is a behaviour that can be influenced by multiple factors and is highly plastic. Recent studies have shown consistency in individual foraging behaviour has serious ecological and evolutionary implications within species and populations. Such information is crucial to understand how species select habitats, and how such selection might allow them to adapt to the environmental changes they face. Five foraging metrics (maximum distance from the colony, bearing from the colony to the most distal point, tortuosity index, total number of dives and mean vectorial dynamic body acceleration were obtained using GPS tracking and accelerometry data in adult Australasian gannets (Morus serrator) from two colonies in southeastern Australia. Individuals were instrumented over two breeding seasons to obtain data to assess factors influencing foraging behaviour and behavioural consistency over multiple timescales (consecutive trips, breeding stages and years) and habitats (pelagic, mixed pelagic and inshore, and inshore). Colony, breeding stage and year were the factors which had the greatest influence on foraging behaviour, followed by sex. Behavioural consistency, measured as the contribution of the individual to the observed variance, was low to moderate for all foraging metrics (0.0-27.05%), with the higher values occurring over shorter timescales. In addition, behavioural consistency was driven by spatio-temporal factors rather than intrinsic characteristics. Behavioural consistency was higher in individuals foraging in inshore than pelagic habitats or mixed pelagic/inshore strategy, supporting suggestions that consistency is favoured in stable environments.

Habitat-specific foraging strategies in Australasian gannets.

Knowledge of top predator foraging adaptability is imperative for predicting their biological response to environmental variability. While seabirds have developed highly specialised techniques to locate prey, little is known about intraspecific variation in foraging strategies with many studies deriving information from uniform oceanic environments. Australasian gannets (Morus serrator) typically forage in continental shelf regions on small schooling prey. The present study used GPS and video data loggers to compare habitat-specific foraging strategies at two sites of contrasting oceanographic regimes (deep water near the continental shelf edge, n=23; shallow inshore embayment, n=26), in south-eastern Australia. Individuals from the continental shelf site exhibited pelagic foraging behaviours typical of gannet species, using local enhancement to locate and feed on small schooling fish; in contrast only 50% of the individuals from the inshore site foraged offshore, displaying the typical pelagic foraging strategy. The remainder adopted a strategy of searching sand banks in shallow inshore waters in the absence of conspecifics and other predators for large, single prey items. Furthermore, of the individuals foraging inshore, 93% were male, indicating that the inshore strategy may be sex-specific. Large inter-colony differences in Australasian gannets suggest strong plasticity in foraging behaviours, essential for adapting to environmental change.

Sexual Size Dimorphism and Body Condition in the Australasian Gannet.

Sexual size dimorphism is widespread throughout seabird taxa and several drivers leading to its evolution have been hypothesised. While the Australasian Gannet (Morus serrator) has previously been considered nominally monomorphic, recent studies have documented sexual segregation in diet and foraging areas, traits often associated with size dimorphism. The present study investigated the sex differences in body mass and structural size of this species at two colonies (Pope's Eye, PE; Point Danger, PD) in northern Bass Strait, south-eastern Australia. Females were found to be 3.1% and 7.3% heavier (2.74 ± 0.03, n = 92; 2.67 ± 0.03 kg, n = 43) than males (2.66 ± 0.03, n = 92; 2.48 ± 0.03 kg, n = 43) at PE and PD, respectively. Females were also larger in wing ulna length (0.8% both colonies) but smaller in bill depth (PE: 2.2%; PD: 1.7%) than males. Despite this dimorphism, a discriminant function provided only mild accuracy in determining sex. A similar degree of dimorphism was also found within breeding pairs, however assortative mating was not apparent at either colony (R2 < 0.04). Using hydrogen isotope dilution, a body condition index was developed from morphometrics to estimate total body fat (TBF) stores, where TBF(%) = 24.43+1.94*(body mass/wing ulna length) - 0.58*tarsus length (r2 = 0.84, n = 15). This index was used to estimate body composition in all sampled individuals. There was no significant difference in TBF(%) between the sexes for any stage of breeding or in any year of the study at either colony suggesting that, despite a greater body mass, females were not in a better condition than males. While the driving mechanism for sexual dimorphism in this species is currently unknown, studies of other Sulids indicate segregation in foraging behaviour, habitat and diet may be a contributing factor.

Combined Use of GPS and Accelerometry Reveals Fine Scale Three-Dimensional Foraging Behaviour in the Short-Tailed Shearwater.

Determining the foraging behaviour of free-ranging marine animals is fundamental for assessing their habitat use and how they may respond to changes in the environment. However, despite recent advances in bio-logging technology, collecting information on both at-sea movement patterns and activity budgets still remains difficult in small pelagic seabird species due to the constraints of instrument size. The short-tailed shearwater, the most abundant seabird species in Australia (ca 23 million individuals), is a highly pelagic procellariiform. Despite its ecological importance to the region, almost nothing is known about its at-sea behaviour, in particular, its foraging activity. Using a combination of GPS and tri-axial accelerometer data-loggers, the fine scale three-dimensional foraging behaviour of 10 breeding individuals from two colonies was investigated. Five at-sea behaviours were identified: (1) resting on water, (2) flapping flight, (3) gliding flight, (4) foraging (i.e., surface foraging and diving events), and (5) taking-off. There were substantial intra- and inter- individual variations in activity patterns, with individuals spending on average 45.8% (range: 17.1-70.0%) of time at sea resting on water and 18.2% (range: 2.3-49.6%) foraging. Individuals made 76.4 ± 65.3 dives (range: 8-237) per foraging trip (mean duration 9.0 ± 1.9 s), with dives also recorded during night-time. With the continued miniaturisation of recording devices, the use of combined data-loggers could provide us with further insights into the foraging behaviour of small procellariiforms, helping to better understand interactions with their prey.

Eating locally: Australasian gannets increase their foraging effort in a restricted range.

During the breeding season, seabirds adopt a central place foraging strategy and are restricted in their foraging range by the fasting ability of their partner/chick and the cost of commuting between the prey resources and the nest. Because of the spatial and temporal variability of marine ecosystems, individuals must adapt their behaviour to increase foraging success within these constraints. The at-sea movements, foraging behaviour and effort of the Australasian gannet (Morus serrator) was determined over three sequential breeding seasons of apparent differing prey abundance to investigate how the species adapts to inter-annual fluctuations in food availability. GPS and tri-axial accelerometer data loggers were used to compare the degree of annual variation within two stages of breeding (incubation and chick rearing) at a small gannet colony situated between two larger, nearby colonies. Interestingly, neither males nor females increased the total distance travelled or duration of foraging trip in any breeding stage (P>0.05 in all cases) despite apparent low prey availability. However, consistently within each breeding stage, mean vectorial dynamic body acceleration (an index of energy expenditure) was greater in years of poorer breeding success (increased by a factor of three to eight), suggesting birds were working harder within their range. Additionally, both males and females increased the proportion of a foraging trip spent foraging in a poorer year across both breeding stages. Individuals from this colony may be limited in their ability to extend their range in years of low prey availability due to competition from conspecifics in nearby colonies and, consequently, increase foraging effort within this restricted foraging area.