Abstract: Aerial flight and breath-hold diving present conflicting morphological and physiological demands, and hence diving seabirds capable of flight are expected to face evolutionary trade-offs regarding locomotory performances. We tested whether Kerguelen shags Phalacrocorax verrucosus, which are remarkable divers, have poor flight capability using newly developed tags that recorded their flight air speed (the first direct measurement for wild birds) with propeller sensors, flight duration, GPS position and depth during foraging trips. Flight air speed (mean 12.7 m s(-1)) was close to the speed that minimizes power requirement, rather than energy expenditure per distance, when existing aerodynamic models were applied. Flights were short (mean 92 s), with a mean summed duration of only 24 min day(-1). Shags sometimes stayed at the sea surface without diving between flights, even on the way back to the colony, and surface durations increased with the preceding flight durations; these observations suggest that shags rested after flights. Our results indicate that their flight performance is physiologically limited, presumably compromised by their great diving capability (max. depth 94 m, duration 306 s) through their morphological adaptations for diving, including large body mass (enabling a large oxygen store), small flight muscles (to allow for large leg muscles for underwater propulsion) and short wings (to decrease air volume in the feathers and hence buoyancy). The compromise between flight and diving, as well as the local bathymetry, shape the three-dimensional foraging range (<26 km horizontally, <94 m vertically) in this bottom-feeding cormorant.

Abstract: Continuous growth, associated with a steady parental food supply, is a general pattern in offspring development. So that young chicks can acquire their locomotor independence, this period is usually marked by a fast maturation of muscles, during which different myosin heavy chain (MyHC) isoforms are expressed. However, parental food provisioning may fluctuate seasonally, and offspring therefore face a challenge to ensure the necessary maturation of their tissues when energy is limited. To address this trade-off we investigated muscle maturation in both the pectoral and pelvic girdles of king penguin chicks. This species has an exceptionally long rearing period (1 year), which is prolonged when parental food provisioning is drastically reduced during the sub-Antarctic winter. Approximately 1 month post hatching, chicks acquire a functional pedestrian locomotion, which uses pelvic muscles, whereas swimming, which uses the pectoral muscles, only occurs 1 year later. We therefore tested the hypothesis that the MyHC content of the leg muscles reaches a mature state before those of the pectoral muscles. We found that leg muscle MyHC composition changed with the progressive acquisition of pedestrian locomotion, whereas pectoral muscle fibres reached their mature MyHC profile as early as hatching. Contrary to our predictions, the acquisition of the adult profile in pectoral muscles could be related to an early maturation of the contractile muscular proteins, presumably associated with early thermoregulatory capacities of chicks, necessary for survival in their cold environment. This differential maturation appears to reconcile both the locomotor and environmental constraints of king penguin chicks during growth.

Abstract: 'Fight-or-flight' stress responses allow animals to cope adaptively to sudden threats by mobilizing energy resources and priming the body for action. Because such responses can be costly and redirect behavior and energy from reproduction to survival, they are likely to be shaped by specific life-history stages, depending on the available energy resources and the commitment to reproduction. Here, we consider how heart rate (HR) responses to acute stressors are affected by the advancing breeding season in a colonial seabird, the king penguin (Aptenodytes patagonicus). We subjected 77 birds (44 males, 33 females) at various stages of incubation and chick-rearing to three experimental stressors (metal sound, distant approach and capture) known to vary both in their intensity and associated risk, and monitored their HR responses. Our results show that HR increase in response to acute stressors was progressively attenuated with the stage of breeding from incubation to chick-rearing. Stress responses did not vary according to nutritional status or seasonal timing (whether breeding was initiated early or late in the season), but were markedly lower during chick-rearing than during incubation. This pattern was obvious for all three stressors. We discuss how 'fight-or-flight' responses may be modulated by considering the energy commitment to breeding, nutritional status and reproductive value of the brood in breeding seabirds.