The effects of season and sex upon the morphology and material properties of keratin in the Svalbard rock ptarmigan (Lagopus muta hyperborea).

The material properties and morphologies of the modified integumentary organs of birds (the keratinous bills, claws and feathers) have evolved to withstand the variety of mechanical stresses imposed by their interaction with the environment. These stresses are likely to vary temporally in seasonal environments and may also differ between the sexes as a result of behavioural dimorphism. Here we investigate the morphology and material properties of the claws of male and female Svalbard ptarmigan (Lagopus muta hyperborea) during the summer and winter using nanoindentation. Despite differences in locomotor demands between the sexes and pronounced seasonal differences in environmental conditions, like ground substrate, ambient temperature and day length, there was no significant difference in Young׳s modulus or hardness between the seasons for each sex. However, when comparing males and females, female claws were significantly harder than those of males and both sexes had significantly wider claws during winter. We propose that wider claws may follow winter claw moulting as the claws are regrown and form an important part of the ptarmigan׳s snowshoe-like foot that is an adaptation to locomotion on snow. Future work focusing on growth rates and more broad measures of material properties in both captive and wild birds is required to determine the extent of seasonal and sex differences in the material properties of their keratinous structures.

Understanding sex differences in the cost of terrestrial locomotion.

Little is known regarding the physiological consequences of the behavioural and morphological differences that result from sexual selection in birds. Male and female Svalbard rock ptarmigans (Lagopus muta hyperborea) exhibit distinctive behavioural differences during the breeding season. In particular, males continuously compete for and defend territories in order to breed successfully, placing large demands on their locomotor system. Here, we demonstrate that male birds have improved locomotor performance compared with females, showing both a lower cost of locomotion (CoL) and a higher top speed. We propose that the observed sex differences in locomotor capability may be due to sexual selection for improved male performance. While the mechanisms underlying these energetic differences are unclear, future studies should be wary when pooling male and female data.

The influence of flight style on the aerodynamic properties of avian wings as fixed lifting surfaces.

The diversity of wing morphologies in birds reflects their variety of flight styles and the associated aerodynamic and inertial requirements. Although the aerodynamics underlying wing morphology can be informed by aeronautical research, important differences exist between planes and birds. In particular, birds operate at lower, transitional Reynolds numbers than do most aircraft. To date, few quantitative studies have investigated the aerodynamic performance of avian wings as fixed lifting surfaces and none have focused upon the differences between wings from different flight style groups. Dried wings from 10 bird species representing three distinct flight style groups were mounted on a force/torque sensor within a wind tunnel in order to test the hypothesis that wing morphologies associated with different flight styles exhibit different aerodynamic properties. Morphological differences manifested primarily as differences in drag rather than lift. Maximum lift coefficients did not differ between groups, whereas minimum drag coefficients were lowest in undulating flyers (Corvids). The lift to drag ratios were lower than in conventional aerofoils and data from free-flying soaring species; particularly in high frequency, flapping flyers (Anseriformes), which do not rely heavily on glide performance. The results illustrate important aerodynamic differences between the wings of different flight style groups that cannot be explained solely by simple wing-shape measures. Taken at face value, the results also suggest that wing-shape is linked principally to changes in aerodynamic drag, but, of course, it is aerodynamics during flapping and not gliding that is likely to be the primary driver.

Seasonal differences in jump performance in the Svalbard rock ptarmigan (Lagopus muta hyperborea).

Fat storage is essential to the survival of many bird species, providing energy reserves, but can have an effect on locomotor performance with an associated potential increase in predation risk. In particular, the ability to initiate flight through jumping is critical to predator avoidance and may be influenced by changes in body mass (Mb). Here we investigate seasonal differences in the jump take-off performance of high Arctic Svalbard rock ptarmigan (Lagopus muta hyperborea) resulting from around a 50% increase in Mb during winter as a result of fat deposition. Using force-plate data and videography, we reveal that, in the absence of alterations to take-off angle, winter Svalbard rock ptarmigan are unable to increase hind-limb power output during jumping to compensate for their increased Mb. As a result, peak take-off velocity is reduced by 42% and jump duration is also extended during winter. The consequences of reduced jumping performance upon Svalbard ptarmigan during winter may be relatively small given their low risk of predation during this season. It may be, however, that the observed reduction in jumping performance when fat may contribute to the sub-maximal pattern of fat acquisition observed in other bird species.