SEASONAL ADAPTATIONS IN THE ENERGETICS AND BIOMECHANICS OF LOCOMOTION IN THE SVALBARD ROCK PTARMIGAN (LAGOPUS MUTA HYPERBOREA)

One of the most striking things about many animals is that they can be defined by the ways in which they move. Moving costs metabolic energy and is a significant contributor to the daily energy balance of organisms and therefore fitness. Balancing energy needs is critically important to species inhabiting areas of limited resources. The metabolic cost of locomotion is influenced by physiological, morphological and behavioural factors that vary across species. The influence of these factors within species is less well understood. The objective of my PhD is to elucidate the potential for variation in locomotor performance, in particular the energy consumed and the biomechanics of locomotion within a species, in response to differences in season, sex, age and the nature of the terrain. The Svalbard ptarmigan (Lagopus muta hyperborea) is the only year-round avian resident of the Arctic archipelago of Svalbard. Svalbard is characterized by extreme photoperiodic and climatic conditions, with 24 hours of daylight in summer and continuous darkness in winter, when ice makes food unpredictable. As a result, ptarmigan annually gain significant fat stores, as much as doubling their body mass in winter. The consequences of such large gains in mass upon the metabolic cost and biomechanics of terrestrial locomotion are yet to be quantified. The Svalbard ptarmigan represents a unique opportunity to gain insight into avian adaptations.Using respirometry, I present evidence that winter birds are able to carry their fat stores at no metabolic cost. Using kinematic and force plate data, I show that acquiring fat results in reduced locomotor performance in terms of speed and take-off ability. As well as exhibiting phenotypic variation, male and female Svalbard ptarmigan are behaviourally very different. I present evidence that these behavioural differences are reflected in the metabolic cost of locomotion. In particular, males are both more efficient and faster than female birds during both summer and winter. I suggest that this results from sexual selection upon male locomotor performance. Furthermore, I present data demonstrating that sub-adult males experiencing their first winter possess the same metabolic and speed capabilities of adults. These data may indicate that selection for improved male locomotor performance may act upon sub-adult birds. Regardless of season, age or sex, Svalbard ptarmigan must locomote on a predominantly sloping terrain. The influence of inclines upon the metabolic cost of locomotion in birds is poorly understood. I provide evidence that at the same degree of incline, the cost of lifting 1 kg by 1 vertical metre is similar regardless of season and is therefore dictated by increased positive work. However, this cost varies according to the degree of incline and may be influenced by gait.The principal findings of the 5 first author papers presented are that behavioural, physiological and morphological variation within a species can have significant impacts upon the metabolic cost of locomotion and other aspects of locomotor performance. The potential for intraspecific differences should therefore be taken into account in future research regarding the patterns of energy expenditure in animals.