Testing foraging optimization models in brown bears: Time for a paradigm shift in nutritional ecology?

How organisms obtain energy to survive and reproduce is fundamental to ecology, yet researchers use theoretical concepts represented by simplified models to estimate diet and predict community interactions. Such simplistic models can sometimes limit our understanding of ecological principles. We used a polyphagous species with a wide distribution, the brown bear (Ursus arctos), to illustrate how disparate theoretical frameworks in ecology can affect conclusions regarding ecological communities. We used stable isotope measurements (δ13 C, δ15 N) from hairs of individually monitored bears in Sweden and Bayesian mixing models to estimate dietary proportions of ants, moose, and three berry species to compare with other brown bear populations. We also developed three hypotheses based on predominant foraging literature, and then compared predicted diets to field estimates. Our three models assumed (1) bears forage to optimize caloric efficiency (optimum foraging model), predicting bears predominately eat berries (~70% of diet) and opportunistically feed on moose (Alces alces) and ants (Formica spp. and Camponotus spp; ~15% each); (2) bears maximize meat intake (maximizing fitness model), predicting a diet of 35%-50% moose, followed by ants (~30%), and berries (~15%); (3) bears forage to optimize macronutrient balance (macronutrient model), predicting a diet of ~22% (dry weight) or 17% metabolizable energy from proteins, with the rest made up of carbohydrates and lipids (~49% and 29% dry matter or 53% and 30% metabolizable energy, respectively). Bears primarily consumed bilberries (Vaccinium myrtillus; 50%-55%), followed by lingonberries (V. vitis-idaea; 22%-30%), crowberries (Empetrum nigrum; 8%-15%), ants (5%-8%), and moose (3%-4%). Dry matter dietary protein was lower than predicted by the maximizing fitness model and the macronutrient balancing model, but protein made up a larger proportion of the metabolizable energy than predicted. While diets most closely resembled predictions from optimal foraging theory, none of the foraging hypotheses fully described the relationship between foraging and ecological niches in brown bears. Acknowledging and broadening models based on foraging theories is more likely to foster novel discoveries and insights into the role of polyphagous species in ecosystems and we encourage this approach.

The cost of research: Lasting effects of capture, surgery and muscle biopsy on brown bear (Ursus arctos) movement and physiology.

Animal models are a key component of translational medicine, helping transfer scientific findings into practical applications for human health. A fundamental principle of research ethics involves weighing the benefits of the research to society against the burden imposed on the animals used for scientific purposes. The utilisation of wild animals for research requires evaluation of the effects of capture and invasive sampling. Determining the severity and duration of these interventions on the animal's physiology and behaviour allows for refining study methodology and for excluding or accounting for biased data. In this study, 39 Scandinavian brown bears (Ursus arctos) captured either while hibernating in winter or via helicopter in summer and that underwent surgery as part of a human health project had their movement, body temperature and timing of onset of hibernation compared with those of 14 control bears that had not been captured during the same period. Bears captured in winter and summer showed decreased movement from den exit until late summer, compared to those in the control group. Bears captured in summer showed reduced movement and body temperature for at least, respectively, 14 and 3 days, with an 11% decrease in hourly distance, compared to pre-capture levels, but did not differ in the timing of hibernation onset. We reveal that brown bear behaviour and physiology can be altered in response to capture and surgery for days to months, post-capture. This has broad implications for the conclusions of wildlife studies that rely upon invasive sampling.