Flipper-bands on penguins: what is the cost of a life-long commitment?

The individual marking of flying and flightless birds has a long history in ornithology. It is the only technique which is cheap, simple and effective, yielding results on bird migration, age-specific annual survival and recruitment. Consequently, hundreds of thousands of birds are annually ringed worldwide. Unfortunately, researchers all too often tend to neglect problems associated with rings and tags. In Antarctic penguins, flipper bands have been used extensively by a variety of nations, and banding is an integral part of the Council for the Conservation of Antarctic Marine Living Resources' (CCAMLR) monitoring programme (Standard method A4). This programme suggests that mortality in penguins wearing bands can be attributed to either (a) prey species availability, (b) predation, (c) weather conditions or (d) other. In this paper, we have attempted to quantify energetic costs associated with wearing a flipper band. For that purpose, freshly caught Adelie penguins (n = 7) were introduced, in Antarctica, into a 21 m long still-water tunnel, where their behaviour and energy consumption were determined via observation and gas respirometry. Birds were either immediately marked with a flipper band and tested in the tunnel for ca 2 h, and then taken out and tested again after removal of the band, or vice-versa. Flipper bands significantly (ANOVA, p = 0.006) increased the power input of Adelie penguins during swimming by 24 % over the speed range of 1.4 to 2.2 m S-', from 17 W kg-' to 21.1 W kg-' (n = 115 and 157 measurements, respectively). The implications of banding on foraging performance and sunival of penguins are discussed. Implantable passive transponders could help overcome such problems

Antennae on transmitters on free-living marine animals: Balancing budgets on the high wire

The effect of externally mounted antennae on the energetics of penguins was studied by mounting various antennae on a transducer fixed to a model Magellanic penguin Spheniscus magellanicus to determine drag, run at speeds of up to 2 m s–1 in a swim canal. For rigid antennae set perpendicular to the water flow, measured drag increased with increasing swim speed. Increasing antenna length (for lengths between 100 and 200 mm) or diameter (for diameters between 1 and 4 mm) resulted in accelerating increased drag as a function of both antenna length and diameter. Where antennae were positioned at acute angles to the water flow, drag was markedly reduced, as was drag at higher speeds in flexible antennae. These results were incorporated in a model on the foraging energetics of free-living Magellanic penguins using data (on swim speeds, intervals between prey encounters, amount ingested per patch and dive durations) derived from previously published work and from a field study conducted on birds from a colony at Punta Norte, Argentina, using data loggers. The field work indicated that free-living birds have a foraging efficiency (net energy gain/net energy loss) of about 2.5. The model predicted that birds equipped with the largest rigid external antennae tested (200 mm × 3 mm diameter), set perpendicular to water flow, increased energy expenditure at normal swim speeds of 1.77 m s–1 by 79% and at prey capture speeds of 2.25 m s–1 by 147%, and ultimately led to a foraging efficiency that was about 5 times less than that of unequipped birds. Highly flexible antennae were shown to reduce this effect considerably. Deleterious antenna-induced effects are predicted to be particularly critical in penguins that have to travel fast to capture prey. Possible measures taken by the birds to increase foraging efficiency could include reduced travelling speed and selection of smaller prey types. Suggestions are made as to how antenna-induced drag might be minimized for future studies on marine diving animal

Krill-feeding behaviour in a chinstrap penguin compared to fish-eating in Magellanic penguins: a pilot study.

Inferring feeding activities from undulations in diving depth profiles is widespread in studies of foraging marine predators. This idea, however, has rarely been tested because of practical difficulties in obtaining an independent estimate of feeding activities at a time scale corresponding to depth changes within a dive. In this study we attempted to relate depth profile undulations and feeding activities during diving in a single Chinstrap Penguin Pygoscelis antarctica, by simultaneously using a conventional time-depth recorder and a recently developed beak-angle sensor. Although failure in device attachments meant that data were obtained successfully from just a part of a single foraging trip, our preliminary results show a linear relationship between the number of depth wiggles and the number of underwater beakopening events during a dive, suggesting that the relative feeding intensity of each dive could be represented by depth-profile data. Underwater beak-opening patterns of this krill-feeding penguin species are compared with recent data from three fish- and squid-feeding Magellanic Penguins Spheniscus magellanicus

Monitoring Penguins at Sea using Data Loggers

The activity of four penguin species at sea was studied using new data loggers. One unit was fixed to the bird's backs and recorded swim speed, swim heading and dive depth from which the three dimensional movements of the birds at sea could be constructed by vector calculations. This unit additionally recorded sea temperature and light intensity. A further, single-channel logger was ingested by the birds and recorded stomach temperature during the periods at sea. Drops in stomach temperature were indicative of prey capture and could be ascribed to specific localities

Lip-reading in remote subjects: An attempt to quantify and separate ingestion, breathing and vocalisation in free-living animals

A new mandibular sensor is presented here based on the use of a Hall sensor, attached to one mandible, opposite a magnet, attached to the other mandible. Changes in sensor voltage, proportional to magnetic field strength, and thus inter-mandibular angle, are recorded in a logger. This system was tested on seven captive Adélie penguins (Pygoscelis adeliae) and three gentoo penguins (Pygoscelis papua) during: (1) feeding trials on land, where birds were given known quantities and types of food; and (2) trials in water where birds were allowed to swim and dive freely. In addition, six free-living Magellanic penguins (Spheniscus magellanicus) were equipped with the system for single foraging trips. Angular signatures were looked for in instances when both captive and free-living birds might open their beaks, and it was discovered that five major behaviours could be identified: ingestion, breathing, calling, head shaking and preening. Captive feeding trials showed that prey mass could be determined with reasonable accuracy (r 2=0.92), and there was some indication that prey type could be resolved if recording frequency were high enough. Vocalisations in Adélie penguins (arc calls) took <0.7 s for mean maximum beak angles of 4.2° (SD 1.3), and were distinguished by their relatively gradual change in beak angle and by their high degree of symmetry. Beak shakings were distinguishable by their short duration (multiple peaks of <0.5 s) and minimal maximum angle (<0.5°). Preening behaviour was apparent due to multiple decreasing peaks (angles <8°). Breathing could be subdivided into that during porpoising, where a characteristic double peak in beak angle was recorded, and that during normal surface rests between dives. During porpoising, only the primary peak (mean maximum beak angle 25.1°, SD 4.7) occurred when the bird was out of the water (mean maximum for second peak 5.9°, SD 4.1). During normal surface rests in free-living birds, breaths could be distinguished as a series of beak openings and closures, showing variation in amplitude and frequency according to an apparent recovery from the previous dive and preparation for the subsequent dive to come. The mandibular measuring system presented shows considerable promise for elucidating many hitherto intractable aspects of the behaviour of free-living animals

To slide or stride: when should Adélie penguins (Pygoscelis adeliae) toboggan?

We noted whether Adélie penguins (Pygoscelis adeliae), when travelling over snow, walked or tobogganed according to gradient, snow friction, or snow penetrability. Both walking and tobogganing penguins reduced stride length and stride frequency, and thus speed, with increasing uphill gradient although tobogganing birds travelled faster and with fewer leg movements. The incidence of tobogganing increased with decreasing friction between penguin and snow. The percentage of penguins tobogganing was also highly positively correlated with increasing snow penetrability. Penguins walking on soft snow must expend additional energy to pull their feet through the snow, whereas tobogganing birds do not sink. It is to be expected that Adélie penguins would utilize the most energetically favourable form of travel which, under almost all conditions, appeared to be tobogganing. Although tobogganing appears to be energetically more efficient than walking, rubbing the feathers over snow increases the coefficient of friction in unpreeened plumage. We propose that a high incidence of tobogganing necessitates increased feather care and that the decision whether to walk or toboggan probably represents a balance between immediate energy expenditure and subsequent energy and time expended maintaining plumage condition