Minke whales maximise energy storage on their feeding grounds

Seasonal trends in energy storage of the minke whale (Balaenoptera acutorostrata), a capital breeder, were investigated in Iceland, a North Atlantic feeding ground. The aim was to better understand the energy acquisition strategies of minke whales and the energetic costs that different reproductive classes face during the breeding season. We modelled total blubber volume, using blubber thickness and morphometric measurements of individual whales. Blubber volume was influenced by body length, and was higher for pregnant females than mature whales. Blubber volume increased linearly through the feeding season at the same rate for mature (mean ± s.e.m.=0.0028±0.00103 m3 day -1; N=61 male, 5 female) and pregnant whales (0.0024±0.00100 m3 day-1; N=49), suggesting that minke whales aim to maximise energy storage while on the feeding grounds. The total amount of blubber accumulated over the feeding season (0.51±0.119 m3 for mature and 0.43±0.112 m3 for pregnant whales), together with energy stored as muscle and intra-abdominal fats, constitutes the total amount of energy available for reproduction (fetus development and lactation) on the breeding grounds, as well as migration, daily field metabolic rates, growth and body maintenance. No seasonal variation was observed for immature whales (N=4 male, 12 female), suggesting that they are investing most of their excess energy into growth rather than reproduction, in order to reach the length of sexual maturity faster and start reproducing earlier. Our novel modelling approach provides insight into large whale bioenergetics and life history strategies, as well as the relationship between single-site measurement of blubber thickness and total blubber volume.

Whale watching disrupts feeding activities of minke whales on a feeding ground

Human disturbances of wildlife, such as tourism, can alter the activities of targeted individuals. Repeated behavioural disruptions can have long-term consequences for individual vital rates (survival and reproduction). To manage these sub-lethal impacts, we need to understand how activity disruptions can influence bioenergetics and ultimately individual vital rates. Empirical studies of the mechanistic links between whale-watching boat exposure and behavioural variation and vital rates are currently lacking for baleen whales (mysticetes). We compared minke whale Balaenoptera acutorostrata behaviour on a feeding ground in the presence and absence of whale-watching boats. Effects on activity states were inferred from changes in movement metric data as well as the occurrence of surface feeding events. Linear mixed effects models and generalised estimation equations were used to investigate the effect of whale-watching boat interactions. Measurement errors were quantified, and their effects on model parameter estimates were investigated using resampling methods. Minke whales responded to whale-watching boats by performing shorter dives and increased sinuous movement. A reduction in the probability of observing longer inter-breath intervals during sinuous movement showed that whale-watching boat interactions reduced foraging activity. Further, the probability of observing surface feeding events also decreased during interactions with whale-watching boats. This indicates that whalewatching boats disrupted the feeding activities of minke whales. Since minke whales are capital breeders, a decrease in feeding success on the feeding grounds due to whale-watching boats could lead to a decrease in energy available for foetus development and nursing on the breeding grounds. Such impact could therefore alter the calving success of this species.

Influence of sea-ice structure on minke whale and other krill-predator distribution

This thesis investigated the extent to which sea-ice structure and complexity could be measured to identify suitable feeding habitat for krill-eating predators such as the Antarctic minke whale (Balaenoptera bonaerensis). The results of this study suggest that the distribution and movement of minke whales is limited by the structure of the sea ice and that the sea ice influences minke whales' habitat use more than their overall distribution. Similar relationships were found for other krill predators such as Adelie penguins and crabeater seals.

Whale distribution in relation to prey abundance and oceanographic processes in the shelf waters of the Western Antarctic Peninsula

The Western Antarctic Peninsula (WAP) is a biologically rich area supporting large standing stocks of krill and top predators (including whales, seals and seabirds). Physical forcing greatly affects productivity, recruitment, survival and distribution of krill in this area. In turn, such interactions are likely to affect the distribution of baleen whales. The Southern Ocean GLOBEC research program aims to explore the relationships and interactions between the environment, krill and predators around Marguerite Bay (WAP) in autumn 2001 and 2002. Bathymetric and environmental variables including acoustic backscattering as an indicator of prey abundance were used to model whale distribution patterns. We used an iterative approach employing (1) classification and regression tree (CART) models to identify oceanographic and ecological variables contributing to variability in humpback Megaptera novaeangliae and minke Balaenoptera acutorstrata whale distribution, and (2) generalized additive models (GAMs) to elucidate functional ecological relationships between these variables and whale distribution. The CART models indicated that the cetacean distribution was tightly coupled with zooplankton acoustic volume backscatter in the upper (25 to 100 m), and middle (100 to 300 m) portions of the water column. Whale distribution was also related to distance from the ice edge and bathymetric slope. The GAMs indicated a persistent, strong, positive relationship between increasing zooplankton volume and whale relative abundance. Furthermore, there was a lower limit for averaged acoustic volume backscatter of zooplankton below which the relationship between whales and prey was not significant. The GAMs also supported an annual relationship between whale distribution, distance from the ice edge and bathymetric slope, suggesting that these are important features for aggregating prey. Our results demonstrate that during the 2 yr study, whales were consistently and predictably associated with the distribution of zooplankton. Thus, humpback and minke whales may be able to locate physical features and oceanographic processes that enhance prey aggregation.

Inferring activity budgets in wild animals to estimate the consequences of disturbances

Activity budgets can provide a direct link to an animal's bioenergetic budget and is thus a valuable unit of measure when assessing human-induced nonlethal effects on wildlife conservation status. However, activity budget inference can be challenging for species that are difficult to observe and require multiple observational variables. Here, we assessed whether whalewatching boat interactions could affect the activity budgets of minke whales (Balaenoptera acutorostrata). We used a stepwise modeling approach to quantitatively record, identify, and assign activity states to continuous behavioral time series data, to estimate activity budgets. First, we used multiple behavioral variables, recorded from continuous visual observations of individual animals, to quantitatively identify and define behavioral types. Activity states were then assigned to each sampling unit, using a combination of hidden and observed states. Three activity states were identified: nonfeeding, foraging, and surface feeding (SF). From the resulting time series of activity states, transition probability matrices were estimated using first-order Markov chains. We then simulated time series of activity states, using Monte Carlo methods based on the transition probability matrices, to obtain activity budgets, accounting for heterogeneity in state duration. Whalewatching interactions reduced the time whales spend foraging and SF, potentially resulting in an overall decrease in energy intake of 42%. This modeling approach thus provides a means to link short-term behavioral changes resulting from human disturbance to potential long-term bioenergetic consequences in animals. It also provides an analytical framework applicable to other species when direct observations of activity states are not possible.

Estimating cumulative exposure of wildlife to non-lethal disturbance using spatially explicit capture-recapture models

Impact assessments often focus on short-term behavioral responses of animals to human disturbance. However, the cumulative effects caused by repeated behavioral disruptions are of management concern because these effects have the potential to influence individuals' survival and reproduction. We need to estimate individual exposure rates to disturbance to determine cumulative effects. We present a new approach to estimate the spatial exposure of minke whales to whalewatching boats in Faxaflõi Bay, Iceland. We used recent advances in spatially explicit capture-recapture modeling to estimate the probability that whales would encounter a disturbance (i.e., whalewatching boat). We obtained spatially explicit individual encounter histories of individually identifiable animals using photo-identification. We divided the study area into 1-km2 grid cells and considered each cell a spatially distinct sampling unit. We used capture history of individuals to model and estimate spatial encounter probabilities of individual minke whales across the study area, accounting for heterogeneity in sampling effort. We inferred the exposure of individual minke whales to whalewatching vessels throughout the feeding season by estimating individual whale encounters with vessels using the whale encounter probabilities and spatially explicit whalewatching intensity in the same area, obtained from recorded whalewatching vessel tracks. We then estimated the cumulative time whales spent with whalewatching boats to assess the biological significance of whalewatching disturbances. The estimated exposure levels to boats varied considerably between individuals because of both temporal and spatial variations in the activity centers of whales and the whalewatching intensity in the area. However, although some whales were repeatedly exposed to whalewatching boats throughout the feeding season, the estimated cumulative time they spent with boats was very low. Although whalewatching boat interactions caused feeding disruptions for the whales, the estimated low cumulative exposure indicated that the whalewatching industry in its current state likely is not having any long-term negative effects on vital rates.

Seasonal variability in whale encounters in the Western Antarctic Peninsula

Cetacean sighting surveys were conducted as part of nine multidisciplinary research cruises over late summer, autumn and winter of 2 years (2001–2003) during the Southern Ocean Global Ocean Ecosystems (SO GLOBEC) program. Sea-ice cover differed markedly between years, with apparent effects on cetacean distribution. No ice was present until late June in 2001, while the previous winter sea ice never fully retreated (>30% cover) during the 2002 or 2003 summer, thus increasing the proportion of thicker and more complex ice, including multi-year floes. Humpback (237 sightings; 537 individuals) and minke (103 sightings: 267 individuals) whales were the most commonly detected species. Data from seven comparable cruises were used to identify habitat for minke and humpback whales over five geographically distinct spatial divisions in the study area. In all years, both species were predominantly found in near coastal habitat, particularly in the fjords where complex habitat likely concentrated prey. In 2002 and 2003 the presence of sea ice provided additional feeding habitat, and the numbers of minkes (in winter) and humpbacks (late summer and autumn) in the area doubled compared with 2001. Humpbacks in particular were concentrated at the ice boundaries during late summer and autumn, while minke numbers increased in the winter that followed and occupied ice-covered areas along the entire shelf edge. Important resource sites for these species are mainly located in near-coastal areas and are used in all years, but when ice margins exist and intersect with resource sites they attract much larger numbers of animals due to the dynamics between sea ice and prey.