Seasonal changes in the diving parameters of king penguins (Aptenodytes patagonicus)

Contrasting conditions at-sea are likely to affect the foraging behaviour of seabirds. However, the effect of season on the dive parameters of penguins is poorly known. We report here on an extensive study of the diving behaviour of king penguins (Aptenodytes patagonicus) over the bird's complete annual cycle at the Crozet Islands. Time-depth recorders were used to record dive duration, bottom duration, post-dive interval, ascent rate and descent rate in breeding adults during different seasons in 1995 and 1996. Seasons included summer (n=6, incubation; n=6, chick brooding), autumn and winter (n=5 and n=3, respectively, chick at the crèche stage), and spring (n=4, birds at the post-moult stage). In all seasons dive duration increased with dive depth, but, for a given depth, dives were longer in winter (6.8 min when averaged over the 100-210 m depth layer) than in spring (4.6 min) and summer (4.4 min). The time spent at the bottom of the dives, which probably represents a substantial part of the feeding time, was much longer in winter (2.5 min per dive for dives over the 100-210 m layer) than during other seasons (1.0-1.4 min), i.e. there was a 2.5-fold augmentation for similar diving depths. Ascent and descent rates increased with increasing dive depth, but no difference in the relationships between rates of ascent and descent and dive depth was found among seasons. Furthermore, for all dive depths, ascent and descent rates were independent of the bottom duration. In all seasons post-dive intervals increased with dive duration and with dive depth, but they were longer in spring (2.3 min for dives over the 100-210 m layer) and summer than in autumn and winter (1.6-1.8 min). The diving efficiency decreased with increasing dive depth and was higher in autumn and winter (0.22-0.29) than in summer and spring (0.15-0.18). The large increase in bottom and dive duration from spring to winter is in agreement with the seasonal drop in prey density, with penguins spending more time searching for prey. In contrast, the consistency of the vertical velocity during contrasting conditions at-sea suggests that the transit time to depth is an important component of the foraging behaviour (scanning of the water column) that is independent of the prey availability. The time budget of the penguins during diving in a fluctuating environment appears to vary primarily during the bottom phase of the dives, with bottom duration increasing with diminishing prey supplies, while post-dive intervals shorten in the same time.     

Foraging strategies and prey encounter rate of free-ranging Little Penguins

There is little information on the effort put into foraging by seabirds, even though it is fundamental to many issues in behavioural ecology. Recent researchers have used changes in the underwater cruising speed of penguins to allude to prey ingestion since accelerations are thought to reflect the encounter and pursuit of prey. In this study, we attached minute accelerometers, to determine flipper beat frequency as a proxy for prey pursuit, to Little Penguins Eudyptula minor foraging in shallow waters in Western Australia. During diving, Little Penguins flapped continuously and at a regular pace of 3.16 Hz while descending the water column and throughout the bottom phase of most dives. However, the frequency and amplitude of wingbeats increased transitorily, reaching 3.5–5.5 Hz, during some dives indicating prey pursuit. Pursuit phases lasted a mean of 2.9±3.3 s and occurred principally during the bottom phases of dives (75.4%). Most dives in all birds (86%) had a clear square-shaped depth profile indicating feeding activity near the seabed in the shallow waters of the bays. Hourly maximum depth, time spent underwater, percentage of dives with pursuit events and catch per unit effort showed an overall increase from zero at ca. 0500 h to a maximum during the hours around mid-day before decreasing to zero by 1900 h. During pursuit phases, Little Penguins headed predominantly downward, probably using the seabed to assist them in trapping their prey. In the light of our results, we discuss depth use by Little Penguins and their allocation of foraging effort and prey capture success as a function of environmental conditions.     

Chinstrap penguins alter foraging and diving behavior in response to the size of their principle prey, Antarctic krill

Penguins may exhibit plasticity in their diving and foraging behaviors in response to changes in prey availability. Chinstrap penguins are dependent predators of Antarctic krill in the Scotia Sea region, but krill populations have fluctuated in recent years. We examined the diet of chinstrap penguins at Livingston Island, South Shetland Islands, in relation to their diving and foraging behavior using time-depth recorders over six breeding seasons: 2002–2007. When krill were smaller, more chinstrap penguins consumed fish. In these years, chinstrap penguins often exhibited a shift to deep dives after sundown, and then resumed a shallower pattern at sunrise. These night dives were unexpectedly deep (up to 110 m) and mean night dive depths sometimes exceeded those from the daytime. The average size of krill in each year was negatively correlated to mean night dive depths and the proportion of foraging trips taken overnight. Based on these patterns, we suggest that when krill were small, penguins increasingly targeted myctophid fish. The average krill size was negatively correlated to the time chinstrap penguins spent foraging which suggests that foraging on smaller krill and fish incurred a cost: more time was spent at sea foraging.     

Comparison of diving behavior and foraging habitat use between chinstrap and gentoo penguins breeding in the South Shetland Islands, Antarctica

Chinstrap, Pygoscelis antarctica, and gentoo, P. papua, penguins are sympatric species that inhabit the Antarctic Peninsula. To evaluate differences in the foraging habitat of these two species, we recorded their foraging locations and diving behavior using recently developed GPS-depth data loggers. The study was conducted on King George Island, Antarctica during the chick-guarding period of both species, from December 2006 to January 2007. The area used for foraging, estimated as the 95% kernel density of dive (>5 m) locations, overlapped partially between the two species (26.4 and 68.5% of the area overlapped for chinstrap and gentoo penguins, respectively). However, the core foraging area, estimated as the 50% kernel density, was mostly separate (12.8 and 25.0% of the area overlapped for chinstrap and gentoo penguins, respectively). Chinstrap penguins tended to use off-shelf (water depth > 200 m) regions (77% of the locations for dives >5 m), whereas gentoo penguins mainly used on-shelf (water depth < 200 m) areas (71% of dive locations). The data on foraging locations, diving behavior, and bathymetry indicated that gentoo penguins often performed benthic dives (28% of dives >5 m), whereas chinstrap penguins almost always used the epipelagic/mid-water layer (96% of dives >5 m). Diving parameters such as diving bottom duration or diving efficiency differed between the species, reflecting differences in the use of foraging habitat. The diving parameters also suggested that the on-shelf benthic layer was profitable foraging habitat for gentoo penguins. Conversely, the relationship between trip duration, date, and stomach content mass suggested that the chinstrap penguins went further from the colony to forage as the season progressed, possibly reflecting a reduction in prey availability near the colony. Our results suggest that chinstrap and gentoo penguins segregated their foraging habitat in the Antarctic coastal marine environment, possibly due to inter- and intra-specific competition for common prey resources.     

Lip-reading in remote subjects: an attempt to quantify and separate ingestion, breathing and vocalisation in free-living animals using penguins as a model

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 (r²=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.     

Geographical variation in the behaviour of a central place forager: Antarctic fur seals foraging in contrasting environments

Foragers show adaptive responses to changes within their environment, and such behavioural plasticity can be a significant driving force in speciation. We investigated how lactating Antarctic fur seals, Arctocephalus gazella, adapt their foraging within two contrasting ecosystems. Location and diving data were collected concurrently, between December 2003 and February 2004, from 43 seals at Bird Island, where krill, Euphausia superba, are the main prey, and 39 at Heard Island, where mostly fish are consumed. Seals at Heard Island were shorter and lighter than those at Bird Island and they spent longer at sea, dived more frequently and spent more time in the bottom phase of dives. Generalized additive mixed effects models showed that diving behaviours differed between the islands. Both populations exploited diel vertically migrating prey species but, on average, Heard Island seals dived deeper and exceeded their estimated aerobic dive limits. We propose that the recovery of the Heard Island population may be limited by the relative inaccessibility and scarcity of food, whereas at Bird Island, the presence of abundant krill resources helps sustain extremely high numbers of seals, even with increased intra- and inter-specific competition. Both populations of fur seals appear to be constrained by their physiological limits, in terms of their optimal diving behaviour. However, there does appear to be some flexibility in strategy at the level of trip with animals adjusting their time at sea and foraging effort, in order to maximize the rate of delivery of energy to their pups.     

Diving behaviour of gentoo penguins,Pygoscelis papua; factors keeping dive profiles in shape

The foraging ecology of seven Gentoo penguins,Pygoscelis papua, breeding at Ardley Island, Antarctica was studied using animal-attached devices which recorded swimming speed, heading and dive depth. Reconstruction of the foraging routes by vectorial analysis of the data indicated that at no time did the birds forage on the sea bed. Swimming speed was relatively constant at 1.7 m s^-1, but rates of descent and ascent in the water column during dives increased with increasing maximum dive depth due to changes in descent and ascent angles. The amount of time spent discending and ascending in the water column increased with maximum dive depth as did the duration spent at the point of maximum depth. Dive profiles were essentially either U-shaped (flat-bottomed dives), or V-shaped (bounce dives). Development of a model based on simple probability theory indicated that the optimal dive profile to maximize the chances of prey acquisition depends on vertical prey distribution and on the visual capabilities of the birds with respect to descent and ascent angles.     

Water column use and forage strategies of female southern elephant seals from Marion Island

The at-sea behaviour of marine top predators provides valuable insights into the distribution of prey species and strategies used by predators to exploit patchily distributed resources. We describe the water column usage and dive strategies of female southern elephant seals from Marion Island tracked between 2004 and 2008. Dives representing increases in forage effort were identified using a method that combines dive type analyses and the calculation of relative amounts of time that animals spend in the bottom phases of dives. Results from this analysis indicate that female elephant seals from Marion Island tend to display lower levels of forage effort closer to the island and display intensive opportunistic forage bouts that occur at a minimum distance of approximately 215 km from the island. Females from Marion Island dived deeper and for longer periods of time, compared to females from other populations. Most animals displayed positive diel vertical migration, evidently foraging pelagically on vertically migrating prey. A few animals displayed periods of reverse (negative) diel vertical migration, however, diving to deeper depths at night, compared to daytime. This behaviour is difficult to explain and prey species targeted during such periods unknown. Our results illustrate plasticity in foraging behaviour of southern elephant seals, as well as inter-population differences in forage strategies.     

Diving behaviour of Little Penguins from four colonies across their whole distribution range: bathymetry affecting diving effort and fledging success

Little Penguins, Eudyptula minor, breed in several small colonies in New Zealand and Australia. In this study, we compare the birds’ diving performances at different sites situated throughout their breeding range. Environmental conditions and breeding success vary drastically amongst colonies, but all birds feed on similar types of prey and face similar limitations on their foraging range. We examined several diving parameters and calculated the proportion of foraging zone available during breeding to examine whether oceanographic and geographic factors in the foraging zone can explain variations in diving behaviour and fledging success among the different colonies. In colonies with high fledging success, Penguin Island and Oamaru, penguins made shallow dives <50 m depth and had lower diving effort. More than 90% of the foraging zone was in waters <50 m depth in these colonies. Motuara Island also has shallow waters with 95% <50 m depth, but the fledging success was low. Phillip Island has only 42% of waters <50 m and comparatively low fledging success. Thus, penguins dived deeper and showed a higher diving effort in colonies with lower fledging success (Motuara Island and Phillip Island), indicating that they were disadvantaged compared to conspecifics from other colonies that dived shallower and with a lesser diving effort. We concluded that bathymetry is an important factor, but not the only one, which influences fledging success.     

How age and sex drive the foraging behaviour in the king penguin

As predicted by life history theory, once recruited into the breeding population and with increasing age, long-lived animals should be able to manage more efficiently the conflict between self-maintenance and reproduction. Consequently, breeding performances should improve with age before stabilizing at a certain level. Using temperature–depth recorders and isotopic analysis, we tested how age affects the foraging behaviour of king penguin Aptenodytes patagonicus during one trip in the chick-rearing phase. Depending on sex and age, king penguins expressed two different foraging strategies. Older birds gained more daily mass per unit body mass than younger ones. Older females conducted shorter trips, dived deeper and performed more prey pursuits. They also had higher blood levels of δ¹⁵N than younger individuals and males indicating sex- and age-specific dietary regimes. However, we found no differences in carbon isotopic signature, suggesting that individuals exploited the same foraging areas independently of sex and age. Our results suggest that king penguins are able to increase the quantity of energy extracted with increasing age and that such a strategy is sex-related. Our study is the first to reveal of an interaction between age and sex in determining foraging efficiency in king penguins.     

Diving behaviour of female northern rockhopper penguins, Eudyptes chrysocome moseleyi, during the brooding period at Amsterdam Island (Southern Indian Ocean)

The pattern and characteristics of diving in 14 female northern rockhopper penguins, Eudyptes chrysocome moseleyi, were studied at Amsterdam Island (37°50′S; 77°31′E) during the guard stage, using electronic time–depth recorders. Twenty-nine foraging trips (27 daily foraging trips and two longer trips including one night) with a total of 16 572 dives of ≥3 m were recorded. Females typically left the colony at dawn and returned in the late afternoon, spending an average of 12 h at sea, during which they performed ∼550 dives. They were essentially inshore foragers (mean estimated foraging range 6 km), and mainly preyed upon the pelagic euphausiid Thysanoessa gregaria, fishes and squid being only minor components of the diet. Mean dive depth, dive duration, and post-dive intervals were 18.4 m (max. depth 109 m), 57 s (max. dive duration 168 s), and 21 s (37% of dive duration), respectively. Descent and ascent rates averaged 1.2 and 1.0 ms⁻¹ and were, together with dive duration, significantly correlated with dive depth. Birds spent 18% of their total diving time in dives reaching 15 to 20 m, and the mean maximum diving efficiency (bottom time:dive cycle duration) occurred for dives reaching 15 to 35 m. The most remarkable feature of diving behaviour in northern rockhopper penguins was the high percentage of time spent diving during daily foraging trips (on average, 69% of their time at sea); this was mainly due to a high dive frequency (∼44 dives per hour), which explained the high total vertical distance travelled during one trip (18 km on average). Diving activity at night was greatly reduced, suggesting that, as other penguins, E. chrysocome moseleyi are essentially diurnal, and locate prey using visual cues. Received: 9 December 1998 / Accepted: 3 March 1999     

Do mountain log skinks (<Emphasis Type="Italic">Pseudemoia entrecasteauxii</Emphasis>) modify their behaviour in the presence of two predators?

Prey often adopt antipredator strategies to reduce the likelihood of predation. In the presence of predators, prey may use antipredator strategies that are effective against a single predator (specific) or that are effective against several predators (nonspecific). Most studies have been confined to single predator environments although prey are often faced with multiple predators. When more than one predator is present, specific antipredator behaviours can conflict and avoidance of one predator may increase vulnerability to another. To test how prey cope with this dilemma, I recorded the behaviours of lizards responding to the nonlethal cues of a bird and snake presented singly and simultaneously. Lizards use specific and conflicting antipredator tactics when confronted with each predator, as evidenced by refuge use. However, when both predators were present, lizards refuge use was the same as in the predator-free environment, indicating that they abandoned refuge use as a primary mechanism for predator avoidance. In the presence of both predators, they reduced their overall movement and time spent thermoregulating. This shift in behaviour may represent a compromise to minimize overall risk, following a change in predator exposure. This provides evidence of plasticity in lizard antipredator behaviour and shows that prey responses to two predators cannot be accurately predicted from what is observed when only one predator is present.Communicated by W. Cooper     

Prey ecology and behaviour affect foraging strategies in the Great Cormorant

The fine link between a particular dive pattern and a specific prey item represents a challenging task in the analysis of marine predator–prey relationships. There is growing evidence that prey type affects diving seabirds’ foraging strategies, dive shapes and underwater activity costs. This study investigates whether a generalist diver, the Great Cormorant Phalacrocorax carbo, modifies the time budget allocated to prey-capture behaviour and breathing strategies (reactive vs. anticipatory) with respect to the prey type (pelagic vs. benthic). Video recordings of 91 Great Cormorants show how the ecology and behaviour of their main prey, Mullets (Mugilidae) and Flounders Platichthys flesus, affect dive/surface durations and the diving pattern. The demersal habit and the low mobility of Flounders leads to an easy access to prey with an anticipatory strategy. Moreover, the patchy distribution of this fish species increases prey-capture rates. Conversely, Mullets exploit the whole water column and are highly mobile, and this is reflected in the need of performing two sequential dives to capture a prey, both longer and likely more expensive, with a consequent switch of strategy from reactive in the searching phase to anticipatory breathing during prey-capture events. This study provides evidence that a generalist diver may switch between different foraging strategies, and it shows how each of them may be optimal under particular ecological conditions. These constraints influence the dynamics that operate within the marine food chains and have relevant implications in managing lagoon areas, including fish ponds.     

Individual boldness affects interspecific interactions in sticklebacks

Within populations of many species, individuals that are otherwise similar to one another in age, size or sex can differ markedly in behaviours such as resource use, risk taking and competitive ability. There has been much research into the implications of such variation for intraspecific interactions, yet little investigation into its role in influencing interspecific interactions outside of a predator–prey context. In this study, we investigated the role of individual-level behavioural variation in determining the outcomes of interactions between two ecologically similar fishes, the threespine and ninespine sticklebacks (Gasterosteus aculeatus and Pungitius pungitius). Experiment 1 asked whether individuals of both species were consistent in their expression of two behaviours: activity in novel surroundings and latency to attack prey. For each behaviour, focal individuals were assayed twice, 10 days apart. Performances were positively correlated between exposures, suggesting behavioural consistency within individuals, at least over this timescale. Experiment 2 revealed not only differences in habitat use described both by species-level variation, with ninespines spending more time in vegetated areas, but also by individual differences, with more active individuals of both species spending more time in open water than in vegetation. Experiment 3 revealed that when heterospecific pairs competed for prey, bolder individuals consumed a greater share, irrespective of species. These findings suggest that individual-level variation can facilitate overlap in habitat use between heterospecifics and also determine the outcomes of resource contests when they meet. We discuss how this might vary between populations as a function of prevailing selection pressures and suggest approaches for testing our predictions.     

Marine aquaculture and bottlenose dolphins’ (<Emphasis Type="Italic">Tursiops truncatus</Emphasis>) social structure

In this study, we investigate association patterns of 249 bottlenose dolphin feeding groups off Sardinia Island (Italy) from January 2000–May 2007 and describe how their association behaviour is related to their response to food patches created by a marine fin fish farm. We also tested the hypothesis that dolphins have different social structures with different feeding activities: Associations should decrease during opportunistic feeding behaviours as it is easier to capture prey, and cooperation is not as necessary. Sixteen individually identified bottlenose dolphins were observed participating in both opportunistic and not opportunistic feeding activities, with a mean of 30 ± 8 times and 9.6 ± 1 times, respectively. Bottlenose dolphins show non-random social behaviour during feeding and this behaviour differs depending on their specific foraging activity. Dolphin associations during feeding can be divided into three categories: acquaintances, affiliates, and feeding associates. Association behaviour during fish farm feeding is consistent with our hypothesis that during opportunistic behaviours, benefits from cooperation decrease, as it is easier to capture prey. Group size homogeneity in both feeding activities demonstrates that the number of dolphins engaging in foraging is not necessarily related with cooperation levels. Moreover, an adult dolphin may prefer to associate with a specific individual, independent of the sex, who shares the same foraging priorities. This study is the first to show how aquaculture is not only directly affecting marine predators but could also indirectly affect their social structure and behaviour.     

In-depth studies of Magellanic penguin (<Emphasis Type="Italic">Spheniscus magellanicus</Emphasis>) foraging: can we estimate prey consumption by perturbations in the dive profile?

A new concept based on analysis of dive depth data was developed to help estimate prey consumption in ten free-ranging Magellanic penguins (Spheniscus magellanicus) that were brooding chicks. By simultaneously analysing the undulations in the dive depth profile (measured by time-depth recorders, TDRs) and beak opening (obtained from the recently developed intra-mandibular angle sensors, IMASEN), it was possible to determine the proportions of the undulations in the dive profile that resulted (or not) in prey capture. This methodology allowed the number of prey consumed to be estimated with a mean error of 10±6% using TDR data alone. If the mean mass of prey is known, then the overall mass of prey consumed per unit time can be determined. Additionally, the method allows estimation of the depth at which prey is taken and thus indicates how penguins exploit the water column. Due to its simplicity, the proposed methodology has applications for other Spheniscus penguin species and should be considered for other marine endotherm divers that show undulations in the dive depth profile.Communicated by O. Kinne, Oldendorf/Luhe     

GPS and time-depth loggers reveal underwater foraging plasticity in a flying diver,the Cape Cormorant

Knowledge on how divers exploit the water column vertically in relation to water depth is crucial to our understanding of their ecology and to their subsequent conservation. However, information is still lacking for the smaller-bodied species, due mostly to size constraints of data-loggers. Here, we report the diving behaviour of a flying diving seabird, the Cape Cormorant Phalacrocorax capensis, weighing 1.0–1.4 kg. Results were obtained by simultaneously deploying small, high resolution and high sampling frequency GPS and time-depth loggers on birds breeding on islands off Western South Africa (34°S, 18°E) in 2008. In all, dive category was assigned to all dives performed by 29 birds. Pelagic dives occurred almost as frequently as benthic dives. Pelagic dives were shallow (mean: 5 m) and took place over seafloors 5–100 m deep. Benthic dives were deeper, occurring on seafloors mainly 10–30 m deep. Dive shape was linked to dive category in only 60% of dives, while the descent rate, ascent rate and bottom duration/dive duration ratio of a dive best explained its dive category. This shows that only the concomitant use of tracking and depth tags can adequately classify diving strategies in a diver like the Cape Cormorant. Diet was mainly Cape Anchovy Engraulis encrasicolis, suggesting that birds probably displayed two contrasted strategies for capturing the same prey. Flexible foraging techniques represent an important key to survival inside the highly productive but heterogeneous Benguela upwelling ecosystem.     

Do activity costs determine foraging tactics for an arctic seabird?

How energy costs affect foraging decisions is poorly understood for marine animals. To provide data relevant to this topic, we examined the relationship between activity levels and foraging behavior by attaching activity recorders to 29 chick-rearing wing-propelled diving birds (thick-billed murres, Uria lomvia) in 1999–2000. We connected the activity during the final dive bout with the prey item we observed being fed to the chicks. After accounting for changes in activity level with depth, activity was highest during the final dive of a dive bout, reflecting maneuvring during prey capture. Pelagic prey items, especially invertebrates (amphipods), were associated with higher depth-corrected activity, leading to shorter dives for a given depth (presumably due to higher oxygen consumption rates) and, thus, shorter search times (lower bottom time for a given depth). Pelagic prey items were likely captured during active pursuit, with the birds actively seeking and pursuing schooling mid-water prey. In contrast, benthic prey involved low activity and extended search times, suggesting that the birds slowly glided along the bottom in search for prey hidden in the sediments or rocks. We concluded that activity levels are important in determining the foraging tactics of marine predators. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The diving behaviour of brooding king penguins (<Emphasis Type="Italic">Aptenodytes patagonicus</Emphasis>) from the Falkland Islands: variation in dive profiles and synchronous underwater swimming provide new insights into their foraging strategies

The diving behaviour of king penguins (Aptenodytes patagonicus) was studied on the Falkland Islands, where a small population (ca. 300 fledglings year^–1) is located at the geographical limit of their breeding range. King penguins rearing newly hatched chicks were equipped with time-depth recorders before leaving for sea. In total, 20,175 dives >3 m were recorded from 12 birds during 15 foraging trips with a mean duration of 5.7±2.3 days. The majority of the trips was directed up to 500 km to the northeast of the breeding colony in slope waters of, and oceanic waters beyond, the Patagonian shelf. Mean time spent underwater accounted for 42±9% of the foraging trip. Mean dive depth achieved was 55±16 m; maximum dive depth recorded was 343 m. Mean dive duration was 159±25 s; maximum dive duration was 480 s. The mean vertical distance covered was 140±65 km trip^–1; and on average birds covered 25 km day^–1. Synchronous diving behaviour was observed in two birds for a period of about 24 h after leaving the colony. Dive depth correlated positively with: (1) light intensity, (2) dive duration and (3) vertical velocities, thus confirming previous findings obtained from conspecifics at other breeding sites and indicating comparable diving behaviour. However, separation of dives according to their profile—V-, U-, or W-shaped—revealed significant differences between certain dive parameters. For a given depth range, bottom time was longer and vertical velocities higher in W-dives than in U-dives. This, together with a higher number of W-dives at dawn and dusk, suggests that foraging is more effective during W-dives than U-dives, and during twilight. These findings imply that king penguins have to make more complex decisions, individually and socially, on the performance of the subsequent dive than previously thought.Communicated by O. Kinne, Oldendorf/Luhe     

Diet of the Humboldt penguin (<Emphasis Type="Italic">Spheniscus humboldti</Emphasis>) in northern and southern Chile

The diet of the Humboldt penguin (Spheniscus humboldti) was examined and compared in two colonies in Chile. Field work was conducted on Pan de Azúcar Island in northern Chile in the breeding season 1998/1999 and on the Puñihuil Islands in southern Chile over two successive breeding seasons during 1997/1998 and 1998/1999. Penguin diet was studied by stomach-pumping birds and analysed by species composition, size and mass of prey. Fish were the dominant prey item at both sites, the contribution of cephalopods and crustaceans varying between sites. The fish prey consisted predominantly of school fish, but there were clear latitudinal differences in fish prey taken. Penguins in the northern colony consumed primarily garfish (Scomberesox saurus), while birds at the southern colony of Puñihuil fed primarily on anchovy (Engraulis ringens), Araucanian herring (Strangomera bentincki) and silverside (Odontesthes regia). The results showed significant differences in terms of numbers of fish taken between the two breeding seasons at Puñihuil. In 1997/1998 penguins consumed almost exclusively anchovy, while they fed primarily on silversides in the successive year. Almost all prey, except stomatopods, were characterised as being pelagic species that occur in relatively inshore water, consistent with the foraging behaviour of Humboldt penguins. The dependence of Humboldt penguins on commercially exploited, schooling prey species makes the species particularly susceptible to changes in prey stocks, due to non-sustainable fisheries management.Communicated by O. Kinne, Oldendorf/Luhe