Research into the terrestrial locomotion of birds is often based upon laboratory treadmill experiments. However, it is unclear how transposable these results are for birds moving in the wild. Here, using video recordings, we compared the kinematics of locomotion (stride frequency, stride length, stance phase, swing phase, duty factor) and speed range of Svalbard rock ptarmigan ( Lagopus muta hyperborea ) under field and laboratory treadmill conditions. Our findings indicate that the kinematics of walking and aerial running are conserved when moving on the treadmill and in the field. Differences, however, were found when grounded running under the two conditions, linked to substrate. Substrate effects were confirmed by analysing trials only moving over very hard snow. In line with laboratory treadmill energetic predictions, wild ptarmigan have a preferred speed during walking and to a lesser extent when aerial running but not when moving with a grounded running gait. The birds were also capable of a higher top speed in the field than that observed during treadmill studies. Our findings demonstrate that laboratory treadmill research provides meaningful information relevant to wild birds while highlighting the importance of understanding the substrate the animals are moving over.

Background: Using Froude numbers (Fr) and relative stride length (stride length: hip height), trackways have been widely used to determine the speed and gait of an animal. This approach, however, is limited by the ability to estimate hip height accurately and by the lack of information related to the substrate properties when the tracks were made, in particular for extinct fauna. By studying the Svalbard ptarmigan moving on snow, we assessed the accuracy of trackway predictions from a species-specific model and two additional Fr based models by ground truthing data extracted from videos as the tracks were being made. Results: The species-specific model accounted for more than 60% of the variability in speed for walking and aerial running, but only accounted for 19% when grounded running, likely due to its stabilizing role while moving faster over a changing substrate. The error in speed estimated was 0-35% for all gaits when using the species-specific model, whereas Fr based estimates produced errors up to 55%. The highest errors were associated with the walking gait. The transition between pendular to bouncing gaits fell close to the estimates using relative stride length described for other extant vertebrates. Conversely, the transition from grounded to aerial running appears to be species specific and highly dependent on posture and substrate. Conclusion: Altogether, this study highlights that using trackways to derive predictions on the locomotor speed and gait, using stride length as the only predictor, are problematic as accurate predictions require information from the animal in question.

Substrate supportiveness is linked to the metabolic cost of locomotion, as it influences the depth to which the foot of a moving animal will sink. As track depth increases animals typically reduce their speed to minimise any potential energetic imbalance. Here we examine how self-selected speed in the Svalbard rock ptarmigan is affected by snow supportiveness and subsequent footprint depth measured using thin-blade penetrometry and 3D photogrammetry, respectively. Our findings indicate that snow supportiveness and footprint depth are poor predictors of speed (r2 = 0.149) and stride length (r2 = 0.106). The ptarmigan in our study rarely sunk to depths beyond the intertarsal joint, regardless of the speed, suggesting that at this relatively shallow depth any increased cost is manageable. 3D reconstructions also indicate that the ptarmigan may exploit the compressive nature of snow to generate thrust during stance, as a trend towards greater foot rotations in deeper footprints was found. It remains unclear if the Svalbard ptarmigan are deliberately avoiding unsupportive snowy substrates. However, if they do, these results would be consistent with the idea that animals should choose routes that minimise energy costs of locomotion. Resumen La firmeza del sustrato se asocial al costo metabólico de la locomoción ya que influencia cuán profundo las extremidades de un animal se hunden al moverse. A medida hundimiento aumenta, usualmente los animales reducen su velocidad para minimizar potenciales desbalances energéticos. En este estudio examinamos cómo la velocidad de la perdiz de la roca de Svalbard es afectada por la firmeza del sustrato y la profundidad de hundimiento de sus patas, usando penetrometría y fotogrametría 3D, respectivamente. Nuestros resultados indican que la firmeza de la nieve y la profundidad de hundimiento de las patas no son buenos predictores de la velocidad (r2 = 0.149) y de la longitud de la zancada (r2 = 0.106). La profundidad de las huellas de las perdices de nuestro estudio rara vez sobrepasó la altura de la articulación intertarsal, independientemente de la velocidad de locomoción, sugiriendo que a profundidades relativamente menores los costos energéticos son manejables. Las reconstrucciones 3D también indican que las perdices podrían aprovechar la naturaleza compresiva de la nieve para generar suficiente empuje durante la fase de soporte, ya que se encontró una tendencia hacia mayores rotaciones de la pata en huellas más profundas. Es incierto si las perdices de Svalbard deliberadamente evitan áreas con nieve más blanda. Sin embargo, si lo hacen, estos resultados serían consistentes con la idea de que los animales deberían seleccionar rutas que minimizan los gastos energéticos en locomoción.

The majority of locomotor research is conducted on treadmills and few studies attempt to understand the differences between this and animals moving in the wild. For example, animals may adjust their gait kinematics or limb posture, to a more compliant limb, to increase stability of locomotion to prevent limb failure or falling on different substrates. Here, using video recordings, we compared locomotor parameters (speed range, stride length, stride frequency, stance duration, swing duration and duty factor) of female Svalbard rock ptarmigan (Lagopus muta hyperborea) moving in the wild over snow to previous treadmill-based research. We also compared the absolute and body size (body mass and limb length)-corrected values of kinematic parameters to published data from males to look for any sex differences across walking and grounded running gaits. Our findings indicate that the kinematics of locomotion are largely conserved between the field and laboratory in that none of the female gaits were drastically affected by moving over snow, except for a prolonged swing phase at very slow walking speeds, likely due to toe dragging. Comparisons between the sexes indicate that the differences observed during a walking gait are likely due to body size. However, sexual dimorphism in body size could not explain the disparate grounded running kinematics of the female and male ptarmigan, which might be linked to a more crouched posture in females. Our findings provide insight into how males and females moving in situ may use different strategies to alleviate the effects of a variable substrate.

Background Using Froude numbers (Fr) and relative stride length (stride length: hip height), trackways have been widely used to determine the speed and gait of an animal. This approach, however, is limited by the ability to estimate hip height accurately and by the lack of information related to the substrate properties when the tracks were made, in particular for extinct fauna. By studying the Svalbard ptarmigan moving on snow, we assessed the accuracy of trackway predictions from a species-specific model and two additional Fr based models by ground truthing data extracted from videos as the tracks were being made. Results The species-specific model accounted for more than 60% of the variability in speed for walking and aerial running, but only accounted for 19% when grounded running, likely due to its stabilizing role while moving faster over a changing substrate. The error in speed estimated was 0-35% for all gaits when using the species-specific model, whereas Fr based estimates produced errors up to 55%. The highest errors were associated with the walking gait. The transition between pendular to bouncing gaits fell close to the estimates using relative stride length described for other extant vertebrates. Conversely, the transition from grounded to aerial running appears to be species specific and highly dependent on posture and substrate. Conclusion Altogether, this study highlights that using trackways to derive predictions on the locomotor speed and gait, using stride length as the only predictor, are problematic as accurate predictions require information from the animal in question.