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.

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.