Scaling of soaring seabirds and its implication for the maximum size of flying pterosaurs

The flight ability of animals is restricted by the scaling effects imposed by physical and physiological factors. In comparisons of the power available from muscle and the mechanical power required to fly, theoretical studies have predicted that the margin between the powers should decrease with body size and that flying animals have a maximum body size. However, predicting an absolute value of this upper limit has been difficult because wing morphology and flight styles vary among species. Albatrosses and petrels have long, narrow, aerodynamically efficient wings and are considered to be soaring birds. Here, using animal-borne accelerometers, we show that scaling analyses of wing-flapping frequencies in these seabirds indicate that the maximum size limit for soaring animals is a body mass of 41 kg and a wingspan of 5.1 m. Soaring seabirds were observed to have two modes of flapping frequencies: vigorous flapping during takeoff and sporadic flapping during cruising flight. In these species, high and low flapping frequencies were found to scale with body mass (_mass_ ^-0.30^ and _mass_ ^-0.18^) in a manner similar to the predictions from biomechanical flight models (_mass_ ^-1/3^ and _mass_ ^-1/6^). The scaling relationships predicted that animals larger than the limit will not be able to flap fast enough to stay aloft under unfavourable wind conditions. Our result therefore casts doubt on the flying ability of large, extinct pterosaurs. The largest extant soarer, the wandering albatross, weighs about 10 kg, which might be a pragmatic limit to maintain a safety margin for sustainable flight and to survive in a variable environment

Diel shifts of swim speeds in king penguins: low swimming speed at night for prey searching and efficient travelling

第6回極域科学シンポジウム[OB] 極域生物圏11月16日（月）　国立極地研究所１階交流アトリウ

Keep the course: travelling penguins consistently orient toward the destination

The Tenth Symposium on Polar Science/Ordinary sessions : [OB] Polar Biology, Wed. 4 Dec. / Entrance Hall (1st floor) , National Institute of Polar Researc

Stable isotopes document the winter foraging ecology of king penguins and highlight connectivity between subantarctic and Antarctic ecosystems

International audienceThe poorly known winter foraging ecology of the king penguin, a major Southern Ocean consumer, was investigated at the subantarctic Crozet Islands where the largest global population breeds. Blood δ13C and δ15N values were used as proxies of the birds’ foraging habitat and diet, respectively, and circulating prolactin levels helped in determining the birds’ reproductive status. Plasma prolactin concentrations showed that king penguin adults of unknown breeding status (n = 52) that were present at the colony in winter were in fact breeders and failed breeders, but were not non -breeders. Circulating prolactin was neither related to δ13C nor δ15N values, thus suggesting that both breeders and failed breeders used the same foraging habitats and fed on the same prey. Plasma and blood cell isotopic values depicted four new relevant biological features on the feeding strategies of king penguins during the critical winter period: (1) 42% of the birds foraged in the distant Antarctic Zone, but 58% fed primarily in subantarctic waters (δ13C), (2) they preyed upon myctophids in both zones (δ15N), (3) individuals were consistent in their foraging strategies over the winter months (δ13C and δ15N), and (4) a higher proportion of females (77%–80%) than males (27%–31%) favored feeding in distant Antarctic waters (δ13C). This study highlights trophic connectivity between subantarctic and Antarctic ecosystems and hence the key role of energy export from Antarctic waters to sustain breeding populations of subantarctic predators, including during the Austral winter

Combined influence of meso-scale circulation and bathymetry on the foraging behaviour of a diving predator, the king penguin (Aptenodytes patagonicus)

Investigating the responses of marine predators to environmental features is of key importance for understanding their foraging behaviour and reproductive success. In this study we examined the foraging behaviour of king penguins breeding at Kerguelen (southern Indian Ocean) in relation to oceanographic and bathymetric features within their foraging ambit. We used ARGOS and Global Positioning System tracking together with Time-Depth-Temperature-Recorders (TDR) to follow the at-sea movements of incubating and brooding king penguins. Combining the penguin behaviour with oceanographic data at the surface through satellite data and at depth through in-situ recordings by the TDRs enabled us to explore how these predators adjusted their horizontal and vertical foraging movements in response to their physical environment. Relating the observed behaviour and oceanographic patterns to local bathymetry lead to a comprehensive picture of the combined influence of bathymetry and meso-scale circulation on the foraging behaviour of king penguins. During both breeding stages king penguins foraged in the area to the south-east of Kerguelen, where they explored an influx of cold waters of southern origin interacting with the Kerguelen Plateau bathymetry. Foraging in the Polar Front and at the thermocline was associated with high prey capture rates. However, foraging trip orientation and water mass utilization suggested that bathymetrically entrained cold-water features provided the most favourable foraging locations. Our study explicitly reports the exploration of bathymetry-related oceanographic features by foraging king penguins. It confirms the presence of Areas of Ecological Significance for marine predators on the Kerguelen Plateau, and suggests the importance of further areas related to the cold-water flow along the shelf break of the Kerguelen Plateau

Can foraging ecology drive the evolution of body size in a diving endotherm?

Within a single animal species, different morphs can allow for differential exploitation of foraging niches between populations, while sexual size dimorphism can provide each sex with access to different resources. Despite being potentially important agents of evolution, resource polymorphisms, and the way they operate in wild populations, remain poorly understood. In this study, we examine how trophic factors can select for different body sizes between populations and sexes in a diving endotherm. Dive depth and duration are positively related to body size in diving birds and mammals, a relationship explained by a lower mass-specific metabolic rate and greater oxygen stores in larger individuals. Based on this allometry, we predict that selection for exploiting resources situated at different depths can drive the evolution of body size in species of diving endotherms at the population and sexual level. To test this prediction, we studied the foraging ecology of Blue-eyed Shags, a group of cormorants with male-biased sexual size dimorphism from across the Southern Ocean. We found that mean body mass and relative difference in body mass between sexes varied by up to 77% and 107% between neighbouring colonies, respectively. Birds from colonies with larger individuals dived deeper than birds from colonies with smaller individuals, when accounting for sex. In parallel, males dived further offshore and deeper than females and the sexual difference in dive depth reflected the level of sexual size dimorphism at each colony. We argue that body size in this group of birds is under intense selection for diving to depths of profitable benthic prey patches and that, locally, sexual niche divergence selection can exaggerate the sexual size dimorphism of Blue-eyed Shags initially set up by sexual selection. Our findings suggest that trophic resources can select for important geographic micro-variability in body size between populations and sexes

Crepuscular foraging by king penguins : behavioral adjustment to the diel vertical migration of prey

The Tenth Symposium on Polar Science/Ordinary sessions : [OB] Polar Biology, Wed. 4 Dec. / Entrance Hall (1st floor) , National Institute of Polar Researc

Scaling of Soaring Seabirds and Implications for Flight Abilities of Giant Pterosaurs

The flight ability of animals is restricted by the scaling effects imposed by physical and physiological factors. In comparisons of the power available from muscle and the mechanical power required to fly, it is predicted that the margin between the powers should decrease with body size and that flying animals have a maximum body size. However, predicting the absolute value of this upper limit has proven difficult because wing morphology and flight styles varies among species. Albatrosses and petrels have long, narrow, aerodynamically efficient wings and are considered soaring birds. Here, using animal-borne accelerometers, we show that soaring seabirds have two modes of flapping frequencies under natural conditions: vigorous flapping during takeoff and sporadic flapping during cruising flight. In these species, high and low flapping frequencies were found to scale with body mass (mass−0.30 and mass−0.18) in a manner similar to the predictions from biomechanical flight models (mass−1/3 and mass−1/6). These scaling relationships predicted that the maximum limits on the body size of soaring animals are a body mass of 41 kg and a wingspan of 5.1 m. Albatross-like animals larger than the limit will not be able to flap fast enough to stay aloft under unfavourable wind conditions. Our result therefore casts doubt on the flying ability of large, extinct pterosaurs. The largest extant soarer, the wandering albatross, weighs about 12 kg, which might be a pragmatic limit to maintain a safety margin for sustainable flight and to survive in a variable environment

Buoyancy under Control: Underwater Locomotor Performance in a Deep Diving Seabird Suggests Respiratory Strategies for Reducing Foraging Effort

BACKGROUND: Because they have air stored in many body compartments, diving seabirds are expected to exhibit efficient behavioural strategies for reducing costs related to buoyancy control. We study the underwater locomotor activity of a deep-diving species from the Cormorant family (Kerguelen shag) and report locomotor adjustments to the change of buoyancy with depth. METHODOLOGY/PRINCIPAL FINDINGS: Using accelerometers, we show that during both the descent and ascent phases of dives, shags modelled their acceleration and stroking activity on the natural variation of buoyancy with depth. For example, during the descent phase, birds increased swim speed with depth. But in parallel, and with a decay constant similar to the one in the equation explaining the decrease of buoyancy with depth, they decreased foot-stroke frequency exponentially, a behaviour that enables birds to reduce oxygen consumption. During ascent, birds also reduced locomotor cost by ascending passively. We considered the depth at which they started gliding as a proxy to their depth of neutral buoyancy. This depth increased with maximum dive depth. As an explanation for this, we propose that shags adjust their buoyancy to depth by varying the amount of respiratory air they dive with. CONCLUSIONS/SIGNIFICANCE: Calculations based on known values of stored body oxygen volumes and on deep-diving metabolic rates in avian divers suggest that the variations of volume of respiratory oxygen associated with a respiration mediated buoyancy control only influence aerobic dive duration moderately. Therefore, we propose that an advantage in cormorants--as in other families of diving seabirds--of respiratory air volume adjustment upon diving could be related less to increasing time of submergence, through an increased volume of body oxygen stores, than to reducing the locomotor costs of buoyancy control