Foraging ecology of Gentoo Penguins Pygoscelis papua at Macquarie Island during the period of chick care

The diet, diving behaviour, swimming velocity and foraging range of Gentoo Penguins Pygoscelis papua were studied at Macquarie Island during the breeding season in the 1993–1994 austral summer. Gentoo Penguins are considered to be inshore feeders, and at Macquarie Island the diet and estimated foraging ranges supported this. The diet consisted of 91.6% fish and 8.3% squid, by mass. The dominant prey taxa were the fish Gymnoscopelus sp. and Paranotothenia magellanica. A mixture of pelagic and benthic prey was consumed, with a greater proportion of benthic species occurring later in the season. The penguins exhibited a strong diurnal pattern in their diving behaviour. Deep diving (≥30 m) began near sunrise (03.00 h) and finished close to sunset (21.00 h). Diving at night was less common and very shallow (<10 m). Early in the breeding season, dive profiles indicated that birds were probably following vertically migrating pelagic prey through the water column and were foraging in waters over 100 m deep. Later in the season, more uniform, shallower depths were used, suggesting an increase in benthic foraging activity. These changes in dive pattern and depth were consistent with the habitat preferences of prey species found in the diet. Gentoo Penguins swam at 1.04 m per s and had a maximum potential foraging range of about 26 km for single-day trips. They tended to forage within 14 km of the colony, with a mean range of 5.4 km. This range encompassed the deep ocean habitat to the west and east of the island and a shallow area to the north.     

Impact of small-scale environmental perturbations on local marine food resources: a case study of a predator,the little penguin

Although the impact of environmental changes on the demographic parameters of top predators is well established, the mechanisms by which populations are affected remain poorly understood. Here, we show that a reduction in the thermal stratification of coastal water masses between 2005 and 2006 was associated with reduced foraging and breeding success of little penguins Eudyptula minor, major bio-indicators of the Bass Strait ecosystem in southern Australia. The foraging patterns of the penguins suggest that their prey disperse widely in poorly stratified waters, leading to reduced foraging efficiency and poor breeding success. Mixed water regimes resulting from storms are currently unusual during the breeding period of these birds, but are expected to become more frequent due to climate change.     

Intra-seasonal variation in foraging behavior among Ad��lie penguins (Pygocelis adeliae) breeding at Cape Hallett, Ross Sea, Antarctica

We investigated intra-seasonal variation in foraging behavior of chick-rearing Adélie penguins, Pygoscelis adeliae, during two consecutive summers at Cape Hallett, northwestern Ross Sea. Although foraging behavior of this species has been extensively studied throughout the broad continental shelf region of the Ross Sea, this is the first study to report foraging behaviors and habitat affiliations among birds occupying continental slope waters. Continental slope habitat supports the greatest abundances of this species throughout its range, but we lack information about how intra-specific competition for prey might affect foraging and at-sea distribution and how these attributes compare with previous Ross Sea studies. Foraging trips increased in both distance and duration as breeding advanced from guard to crèche stage, but foraging dive depth, dive rates, and vertical dive distances travelled per hour decreased. Consistent with previous studies within slope habitats elsewhere in Antarctic waters, Antarctic krill (Euphausia superba) dominated chick meal composition, but fish increased four-fold from guard to crèche stages. Foraging-, focal-, and core areas all doubled during the crèche stage as individuals shifted distribution in a southeasterly direction away from the coast while simultaneously becoming more widely dispersed (i.e., less spatial overlap among individuals). Intra-specific competition for prey among Adélie penguins appears to influence foraging behavior of this species, even in food webs dominated by Antarctic krill.     

Flying south: Foraging locations of the Hutton's shearwater (Puffinus huttoni) revealed by Time‐Depth Recorders and GPS tracking

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Into the deep: the functionality of mesopelagic excursions by an oceanic apex predator

Comprehension of ecological processes in marine animals requires information regarding dynamic vertical habitat use. While many pelagic predators primarily associate with epipelagic waters, some species routinely dive beyond the deep scattering layer. Actuation for exploiting these aphotic habitats remains largely unknown. Recent telemetry data from oceanic whitetip sharks (Carcharhinus longimanus) in the Atlantic show a strong association with warm waters (>20°C) less than 200 m. Yet, individuals regularly exhibit excursions into the meso‐ and bathypelagic zone. In order to examine deep‐diving behavior in oceanic whitetip sharks, we physically recovered 16 pop‐up satellite archival tags and analyzed the high‐resolution depth and temperature data. Diving behavior was evaluated in the context of plausible functional behavior hypotheses including interactive behaviors, energy conservation, thermoregulation, navigation, and foraging. Mesopelagic excursions (n = 610) occurred throughout the entire migratory circuit in all individuals, with no indication of site specificity. Six depth‐versus‐time descent and ascent profiles were identified. Descent profile shapes showed little association with examined environmental variables. Contrastingly, ascent profile shapes were related to environmental factors and appear to represent unique behavioral responses to abiotic conditions present at the dive apex. However, environmental conditions may not be the sole factors influencing ascents, as ascent mode may be linked to intentional behaviors. While dive functionality remains unconfirmed, our study suggests that mesopelagic excursions relate to active foraging behavior or navigation. Dive timing, prey constituents, and dive shape support foraging as the most viable hypothesis for mesopelagic excursions, indicating that the oceanic whitetip shark may regularly survey extreme environments (deep depths, low temperatures) as a foraging strategy. At the apex of these deep‐water excursions, sharks exhibit a variable behavioral response, perhaps, indicating the presence or absence of prey.     

Group Size in Foraging African Penguins (Spheniscus demersus)

Diving synchrony was examined for varying group sizes of African penguins (Spheniscus demersus) travelling to their foraging grounds from their breeding islands. Groups of fewer than 12 birds always dived synchronously, whereas groups of more than 17 birds always dived asynchronously. Since travelling penguins do not dive deeply, large groups of birds can remain together irrespective of diving synchronization. Observations from boats showed that foraging penguins rarely occurred in groups of more than 17 birds. We suggest that groups of penguins that do not have synchronized dives cannot forage effectively, because foraging penguins dive deeply.     

Dive bout organization in the Chinstrap penguin at seal island, antarctica

Diving behavior of 2 breeding Chinstrap penguins (Pygoscelis antarctica) was studied focusing first and primarily on dive bouts rather than dives themselves. Analysis of dive bout organization revealed (1) though there are differences between solitary dives and dive bouts in dive duration and dive depth, the first dives of dive bouts do not differ from solitary dives in the dive parameters, (2) mean dive duration during bout correlates positively to both mean dive depth during bout and mean surface interval during bout, while number of dives during bout negatively correlates to both cost (consumed energy) and duration of a dive cycle during bout. These findings suggest the following possibilities on foraging behavior of penguins: (1) their decision to repeat diving depends on the result of the first dive at a site, and the first dives of bouts would tend to be searching or evaluating dives though they would be also successful foraging dives, (2) they repeat diving at a foraging patch until foraging efficiency decrease to a threshold of diminishing returns.     

Time/depth usage of Adélie penguins: an approach based on dive angles

Data on the swim speed, dive depth and feeding rates of three Adélie penguins (Pygoscelis adeliae) foraging in summer 1998/1999 in Adélie Land, Antarctica were collected using dorsally-mounted loggers, in tandem with oesophageal temperature sensors. Swim speed could be integrated, together with the rate of change of depth, to determine dive and return-to-surface angles. Overall, birds increased rates of change of depth during commuting phases so that dive angles were steeper in dives terminating at greater depths. Angles of descent and ascent during feeding dives were greater than during non-feeding dives. Variation in the descent angle over time of particular dives was generally less than 10°, but the angles of the ascent phases varied more widely. The importance of selecting the optimum descent and ascent angles with respect to prey exploitation, oxygen stores and time gained in the feeding area over the course of a dive by diving at a steeper angle is discussed.     

Water temperature sampling by foraging Brünnich's Guillemots with bird-borne data loggers

We describe the features of waters where seabirds were feeding by sampling vertical water temperature profiles with data loggers mounted on five Brünnich's Guillemots in Svalbard, Norway. The guillemots foraged in a cold water (−0.5–0.5°C SST (sea surface temperature)) by making 1.8 dive bouts in short trips (32–257 min duration) as well as in moderate (0.5–2.0°C SST) and warm waters (2.5–4.0°C SST) by making 6.0 dive bouts during long trips (411–688 min duration). Judging from outbound flying time (15.7–24.4 min), time between dive bouts (23.9–43.3 min) and water types, the birds probably fed in fjord or coastal waters during short trips and in both coastal and offshore waters during long trips. Water temperature and diving behaviour can be simultaneously recorded by small data loggers, which therefore will provide useful information on marine features and foraging activity of top predators.     

Diving behaviour and energetics in breeding little penguins (Eudyptula minor)

We present data on the diving behaviour and the energetics of breeding little penguins in Tasmania, Australia. Using an 18 m long still water canal in conjunction with respirometry, we determined the energy requirements while diving. Using electronic devices measuring dive depth or swimming speed, we investigated the foraging behaviour at sea. Cost of Transport was calculated to be minimal at the speed the birds prefer at sea (1.8 m/s) and averaged 11.1 J/kg/m (power requirements at that speed: 20.0 W/kg). Metabolic rate of little penguins resting in water was found to be 8.5 W/kg. The externally-attached devices had no significant influence on the energy expenditure.
Foraging trips can be divided into four distinct phases with different diving behaviours. A mean of 500 dives was executed per foraging trip lasting about 18 hours with 60% of this time being spent swimming. The total distance travelled averaged 73 km per day, although foraging range was about 12km. Mean swimming speed of little penguins at sea was 1.8 m/s, maximum swimming speed was 3.3 m/s. More than 50% of all dives had maxima not exceeding 2 m. Maximum depth reached was 27 m. Mean dive duration was 21 s. There were inter-sex differences in diving behaviour as well as changes in foraging behaviour over the breeding period. Aerobic dive limits (ADL) in the wild were estimated between 42 and 50 s. From the swim canal experiments we derived an ADL of 44 s. Total oxygen stores were calculated to be 45 ml O₂/kg. Only 2% of all dives exceeded the ADL. FMRs at sea were calculated to be between 1280 and 1500 kJ/kg/d according to chick size. The yearly food requirements of a breeding little penguin amount to 114 kg.     

Importance of the large copepod Paraeuchaeta antarctica (Giesbrecht, 1902) in coastal waters and the diet of seabirds at Kerguelen, Southern Ocean

The importance of the euchaetid copepod Paraeuchaeta antarcticain the subantarctic pelagic ecosystem was quantified in thecoastal waters of the Golfe du Morbihan at Kerguelen Islandsby comparing food samples from two diving seabirds with concurrentnet samples taken within the predator foraging area. Paraeuchaetaantarctica occurred in very high densities (up to 30 individualsm^-3 and 96 mg dry weight m^-3) in the water column, being moreabundant in the deepest part of the gulf than in shallow watersor at the more offshore shelf stations. The common diving petrelfeeds almost exclusively on crustaceans, its diet being dominatedby the amphipod Themisto gaudichaudii (52% by number and 84%by reconstituted mass) and P. antarctica (33% and 16%, respectively).Rockhopper penguins preyed upon crustaceans and fish, with fourtaxa being important, namely T. gaudichaudii (37% and 23%, respectively),Euphausia vallentini (24% and 41%), postlarval fish (10% and24%), and P. antarctica (13% and 3%). Paraeuchaeta antarcticadominated numerically in 21% of the diving-petrel food samplesand in 12% of penguin samples. The two bird species segregatedby preying upon different developmental stages of P. antarctica,diving petrels fed equally on CV of both sexes and CVI, whilepenguins fed on CVI only. Comparison of P. antarctica foundin net and food samples indicated no prey selection by commondiving petrels that caught the different copepod stages in proportionto their availability in the water column. On the other hand,the diving performance of penguins, which is better than thatof the petrels (mean maximum dive depths 69 m for penguins versus32 m in petrels), allows them to catch CVI in deeper waters,probably near the bottom. Our study shows that P. antarcticais a major component of the coastal macrozooplankton communityand a significant prey for two species of diving seabirds inhabitingKerguelen. This is also the first record of a copepod speciesas a prey for penguin, and the first to highlight P. antarcticain the food of austral seabirds.     

Annual and seasonal changes in foraging site and diving behavior in Adélie penguins

Foraging sites, diet, and diving behavior of chick-rearing Adélie penguins, Pygoscelis adeliae, in fast sea-ice areas were investigated during two consecutive seasons with contrasting sea-ice conditions. During 1995/1996, fast sea ice covered the foraging range of penguins during the whole breeding season. In contrast, during 1996/1997, sea ice covered the area in December 1996, but gradually thinned and finally broke up, so that open sea appeared along the coast during February 1997. Foraging sites were concentrated in a small area in 1995/1996 and spread over a wider area in 1996/1997 as more small open-water areas were available. In both seasons, parents traveled to more distant foraging sites as the season progressed and, consequently, the foraging-trip duration increased. In both years, Euphausia superba and Pagothenia borchgrevinki dominated the diet in the early part of the season, while later in the season penguins fed mainly on E. superba in 1995/1996 and Pagothenia borchgrevinki and E. crystallorophias in 1996/1997. In 1995/1996, penguins tended to dive deeper—albeit for a relatively shorter duration—when feeding mainly on krill compared to when feeding on fish. In 1996/1997, there was no difference in the dive depth and duration between krill- and fish-eating trips. Our results suggest that prey distribution changes annually and seasonally, probably according to sea-ice conditions, and that consequently penguins modify their foraging sites, diving patterns, and diet according to these changes.     

Tufted ducks Aythya fuligula do not control buoyancy during diving

Work against buoyancy during submergence is a large component of the energy costs for shallow diving ducks. For penguins, buoyancy is less of a problem, however they still seem to trade‐off levels of oxygen stores against the costs and benefits of buoyant force during descent and ascent. This trade‐off is presumably achieved by increasing air sac volume and hence pre‐dive buoyancy (B_pre) when diving deeper. Tufted ducks, Aythya fuligula, almost always dive with nearly full oxygen stores so these cannot be increased. However, the high natural buoyancy of tufted ducks guarantees a passive ascent, so they might be expected to decrease B_pre before particularly deep, long dives to reduce the energy costs of diving. Body heat lost to the water can also be a cause of substantial energy expenditure during a dive, both through dissipation to the ambient environment and through the heating of ingested food and water. Thus dive depth (d_d), duration and food type can influence how much heat energy is lost during a dive. The present study investigated the relationship between certain physiological and behavioural adjustments by tufted ducks to d_d and food type. Changes in B_pre, deep body temperature (T_b) and dive time budgeting of four ducks were measured when diving to two different depths (1.5 and 5.7 m), and for two types of food (mussels and mealworms). The hypothesis was that in tufted ducks, B_pre decreases as d_d increases. The ducks did not change B_pre in response to different diving depths, and thus the hypothesis was rejected. T_b was largely unaffected by dives to either depth. However, diving behaviour changed at the greater d_d, including an increase in dive duration and vertical descent speed. Behaviour also changed depending on the food type, including an increase in foraging duration and vertical descent speed when mussels were present. Behavioural changes seem to represent the major adjustment made by tufted ducks in response to changes in their diving environment.     

Diving pattern and performance in relation to foraging ecology in the gentoo penguin, Pygoscelis papua

We present data on diving pattern and performance (dive depth, duration, frequency and organization during the foraging trip) in gentoo penguins Pygoscelis papua , obtained using time-depth recorders ( n = 9 birds, 99 foraging trips). These data are used to estimate various parameters of foraging activity, e.g. foraging range, prey capture rates, and are compared in relation to breeding chronology. Foraging trip duration was 6 h and 10 h, and trip frequency 1.0/day and 0.96/day, during the brooding and creche periods, respectively. Birds spent on average 52%of each foraging trip diving. Dive depth and duration were highly bimodal: shallow dives (< 21 m) averaged 4 m and 0.23 min, and deep dives (> 30 m) 80 m and 2.5 min, respectively. Birds spent on average 71%and 25%of total diving time in deep and shallow dives, respectively. For deep dives, dive duration exceeded the subsequent surface interval, but shallow dives were followed by surface intervals 2–3 times dive duration. We suggest that most shallow dives are searching/exploratory dives and most deep dives are feeding dives. Deep dives showed clear diel patterns averaging 40 m at dawn and dusk and 80–90 m at midday. Estimated foraging ranges were 2.3 km and 4.1 km during the brood and creche period, respectively. Foraging trip duration increased by 4 h between the brood and creche periods but total time spent in deep dives (i.e. time spent feeding) was the same (3 h). Of 99 foraging trips, 56%consisted of only one dive bout and 44%of 2–4 bouts delimited by extended surface intervals > 10 min. We suggest that this pattern of diving activity reflects variation in spatial distribution of prey rather than the effect of physiological constraints on diving ability.     

At-Sea Associations in Foraging Little Penguins

Prey distribution, patch size, and the presence of conspecifics are important factors influencing a predator’s feeding tactics, including the decision to feed individually or socially. Little is known about group behaviour in seabirds as they spend most of their lives in the marine environment where it is difficult to observe their foraging activities. In this study, we report on at-sea foraging associations of little penguins (Eudyptula minor) during the breeding season. Individuals could be categorised as (1) not associating; (2) associating when departing from and/or returning to the colony; or (3) at sea when travelling, diving or performing synchronised dives. Out of 84 separate foraging tracks, 58 (69.0%) involved associations with conspecifics. Furthermore, in a total of 39 (46.4%), individuals were found to dive during association and in 32 (38.1%), individuals were found to exhibit synchronous diving. These behaviours suggest little penguins forage in groups, could synchronise their underwater movements and potentially cooperate to concentrate their small schooling prey.     

Does Foraging Performance Change with Age in Female Little Penguins (Eudyptula minor)?

Age-related changes in breeding performance are likely to be mediated through changes in parental foraging performance. We investigated the relationship of foraging performance with age in female little penguins at Phillip Island, Australia, during the guard phase of the 2005 breeding season. Foraging parameters were recorded with accelerometers for birds grouped into three age-classes: (1) young, (2) middle age and (3) old females. We found the diving behaviour of middle-aged birds differed from young and old birds. The dive duration of middle age females was shorter than that of young and old birds while their dive effort (measure for dive and post-dive duration relation) was lower than that of young ones, suggesting middle-aged birds were in better physical condition than other ones. There was no difference in prey pursuit frequency or duration between age classes, but in the hunting tactic. Females pursued more prey around and after reaching the maximum depth of dives the more experienced they were (old > middle age > young), an energy saving hunting tactic by probably taking advantage of up-thrust momentum. We suggest middle age penguins forage better than young or old ones because good physical condition and foraging experience could act simultaneously.     

Foraging behavior of Adélie penguins during incubation period in Lützow-Holm Bay

The diving behavior of Adélie penguins Pygoscelis adeliae was investigated using time–depth recorders during the incubation period in the fast sea-ice area of Lützow-Holm Bay, Antarctica. Dive profiles and activity/time allocation suggested that penguins were obligated to walk on the fast-ice for 90–100 km until a polynya, which they used as an access to the pack-ice zone. Dive depth did not differ between males and females, though males’ dive duration was longer than that of females. Dive depth was slightly shallower and dive duration was shorter during the incubation than during the chick-rearing phase. Birds dove throughout the day, although less frequently around midnight, and there was no clear diel change in dive depth. This daily dive pattern during incubation period was similar to that previously observed during the chick-rearing period in a fast sea-ice area, but differed from that observed in sea-ice-free area. Variations in diving behavior resulted from different environmental conditions, such as foraging area with different sea-ice condition, as well as from different life history strategies.     

Foraging behaviour of Magellanic Penguins during the early chick‐rearing period at Isla de los Estados,Argentina

Studies of the at‐sea distribution and trophic ecology of penguins are essential to understand their role in the broader marine food web. Magellanic Penguins Spheniscus magellanicus have a wide distribution and their foraging behaviour varies across breeding sites and between sexes, among others. In this study, we characterized the at‐sea areas, the diving strategies and the relative trophic level of Magellanic Penguins breeding at Isla de los Estados, Argentina, during the early chick‐rearing period. In addition, we quantified the interannual, sexual and individual variability in those parameters during three breeding seasons (2011–2013) using devices recording position and dive depth, and obtained blood samples for stable isotope analysis. During the early chick‐rearing period, Magellanic Penguins showed small differences between the sexes in their foraging behaviour and large overlap in the at‐sea areas used, suggesting no intraspecific variation between sexes. Although there was interannual variability in the foraging behaviour and the trophic level of the penguins, most of the studied nests managed successfully to raise both chicks during the first stage of the breeding cycle (guard stage). The foraging ecology of Magellanic Penguins from this colony was comparable with results of past studies at other breeding colonies. This study contributes to the identification of important at‐sea areas for Magellanic Penguins at the southern edge of their distribution and also to the identification of possible threats in the study area such as interaction with fisheries.     

Diel and lunar variation in diving behavior of rough-toothed dolphins (Steno bredanensis) off Kauaʻi,Hawaiʻi

Observational studies describe rough-toothed dolphins (Steno bredanensis) actively foraging during the day on epipelagic species. Using data from depth-transmitting satellite tags deployed on nine individuals off Kauaʻi, we investigated diving behavior and the effects of lunar phase and solar light levels on vertical movements. Overall, tagged rough-toothed dolphins primarily used near-surface waters, spending between 83.6% and 93.7% of their time in the top 30 m of the water column. When diving, grand mean, median, and maximum dive depths were 76.9 m, 67.5 m, and 399.5 m, although individuals were in water with depths from approximately 700–1,450 m. Dive rates varied by time of day, being lowest during the day and at dawn and highest at dusk and night. Dives were deepest (M = 133.7 m, SD = 52.6 m, median = 106.5 m) and longest (M = 4.0 min, SD = 0.4 min, median = 4.0 min) at dusk, suggesting dolphins were taking advantage of prey rising to the surface in response to reduced light levels. Lunar phase indirectly affected diving, with deeper and longer dives occurring with increasing illumination. The variations in dive behavior across solar and lunar cycles indicate diving patterns shift based on the distribution of prey.     

Diving and foraging behaviour of Adélie penguins in areas with and without fast sea-ice

The diving and foraging behaviours of Adélie penguins, Pygoscelis adeliae, rearing chiks at Hukuro Cove, Lützow-Holm Bay, where the fast sea-ice remained throughout summer, were compared to those of penguins at Magnetic Island, Prydz Bay, where the fast sea-ice disappeared in early January. Parent penguins at Hukuro Cove made shallower (7.1–11.3 m) but longer (90–111 s) dives than those at Magnetic Island (22.9 m and 62 s). Dive duration correlated with dive depth at both colonies (r ² = 0.001 ∼ 0.90), but the penguins atg Hukuro Cove made longer dives for a given depth. Parents at Hukuro Cove made shorter foraging trips (8.1–14.4 h) with proportionally longer walking/swimming (diving < 1 m) travel time (27–40% of trip duration) and returned with smaller meals (253–293 g) than those at Magnetic Island, which foraged on average for 57.2 h, spent 2% of time walking/swimming ( < 1 m) travel, and with meals averaging 525 g. Trip duration at both colonies correlated to the total time spent diving. Trip duration at Hukuro Cove, but not at Magnetic Island, increased as walking/swimming ( < 1 m) travel time increased. These differences in foraging behaviour between colonies probably reflected differences in sea-ice cover and the availability of foraging sites. Received: 3 November 1995/Accepted: 29 May 1996