Foraging strategies of incubating and brooding king penguins Aptenodytes patagonicus

For oceanic birds like king penguins, a major constraint is the separation of foraging areas from the breeding colony, largely because swimming increases foraging costs. However, the relationship between foraging strategy and breeding stage has been poorly investigated. Using time-depth recorders, we studied the diving behaviour of two groups of king penguins that were either incubating or brooding chicks at Crozet Islands (Southern Indian Ocean) at the same period of the year. Although birds with chicks had the highest predicted energy demand, they made foraging trips half as long as incubating birds (6 vs. 14 days) and modified their time and depth utilisation. Birds with chicks dived deeper during daylight (mean maximum depth of 280 m vs. 205 m for those incubating). At night, birds with chicks spent twice as much time diving as those incubating, but birds at both stages never dived beyond 30 m. Movements to greater depths by brooding birds are consistent with the vertical distribution of myctophid fish which are the main prey. As chick provisioning limits trip duration, it is suggested that it is more efficient for parents to change their diving patterns rather than to restrict their foraging range. Received: 23 June 1997 / Accepted: 3 November 1997     

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.     

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.     

Activity Time Budget during Foraging Trips of Emperor Penguins

We developed an automated method using depth and one axis of body acceleration data recorded by animal-borne data loggers to identify activities of penguins over long-term deployments. Using this technique, we evaluated the activity time budget of emperor penguins (n = 10) both in water and on sea ice during foraging trips in chick-rearing season. During the foraging trips, emperor penguins alternated dive bouts (4.8±4.5 h) and rest periods on sea ice (2.5±2.3 h). After recorder deployment and release near the colony, the birds spent 17.9±8.4% of their time traveling until they reached the ice edge. Once at the ice edge, they stayed there more than 4 hours before the first dive. After the first dive, the mean proportions of time spent on the ice and in water were 30.8±7.4% and 69.2±7.4%, respectively. When in the water, they spent 67.9±3.1% of time making dives deeper than 5 m. Dive activity had no typical diurnal pattern for individual birds. While in the water between dives, the birds had short resting periods (1.2±1.7 min) and periods of swimming at depths shallower than 5 m (0.25±0.38 min). When the birds were on the ice, they primarily used time for resting (90.3±4.1% of time) and spent only 9.7±4.1% of time traveling. Thus, it appears that, during foraging trips at sea, emperor penguins traveled during dives >5 m depth, and that sea ice was primarily used for resting. Sea ice probably provides refuge from natural predators such as leopard seals. We also suggest that 24 hours of sunlight and the cycling of dive bouts with short rest periods on sea ice allow emperor penguins to dive continuously throughout the day during foraging trips to sea.     

Maximum diving depths of northern rockhopper penguins (Eudyptes chrysocome moseleyi) at Amsterdam Island

 The mean maximum dive depth from 49 foraging bouts by northern rockhopper penguins, measured using capillary-tube depth gauges, was 66±4 m (12–168 m). There were no differences in the maximum dive depths between male and female penguins. Northern rockhopper penguins dived deeper in early than in late creche stages (83±7 vs 57±4 m), and this was associated with probable dietary changes, squid dominating the diet by mass (44%) in November, and fish (64%) in December 1994 at Amsterdam Island. Received: 10 January 1996/Accepted: 31 March 1996     

Magellanic Penguins Spheniscus magellanicus commuting through San Julian Bay; do current trends induce tidal tactics?

The behaviour of Magellanic Penguins Spheniscus magellanicus commuting between their foraging areas and breeding areas in San Julian Bay, Argentina was studied to examine whether tidal rip currents affected travelling patterns. Although there was no apparent relationship between departure and arrival patterns and the state of the tidal cycle, birds travelling against the current dived for longer periods and had shorter rests on the surface than birds travelling with the current. In addition, birds swimming against the current hugged the banks of tidal rivers much more closely than did birds swimming with the current, thus reducing the magnitude of the current against which they had to swim. In cases of extremely high current speeds, birds travelling upstream walked. Models regarding the energetics of movement indicate that the strategies adopted by Magellanic Penguins can result in substantial energy savings.     

Diving depths of Shags Phalacrocorax aristotelis breeding on the Isle of May

Information on maximum dive depth and the time spent at various depths was obtained from 49 Shags Phalacrocorax aristotelis. On average, the maximum dive depth was 33–35 m; the overall maximum was 43 m. Shags dived repeatedly to the same depth and spent c. 55% of the time between 25 and 34 m which indicated that they were foraging close to the seabed. About 46% of the time underwater was spent foraging and 54% travelling. Average underwater swimming speed was 1.7-1.9 m per second.     

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.     

Straight Line Foraging in Yellow-Eyed Penguins: New Insights into Cascading Fisheries Effects and Orientation Capabilities of Marine Predators

Free-ranging marine predators rarely search for prey along straight lines because dynamic ocean processes usually require complex search strategies. If linear movement patterns occur they are usually associated with travelling events or migratory behaviour. However, recent fine scale tracking of flying seabirds has revealed straight-line movements while birds followed fishing vessels. Unlike flying seabirds, penguins are not known to target and follow fishing vessels. Yet yellow-eyed penguins from New Zealand often exhibit directed movement patterns while searching for prey at the seafloor, a behaviour that seems to contradict common movement ecology theories. While deploying GPS dive loggers on yellow-eyed penguins from the Otago Peninsula we found that the birds frequently followed straight lines for several kilometres with little horizontal deviation. In several cases individuals swam up and down the same line, while some of the lines were followed by more than one individual. Using a remote operated vehicle (ROV) we found a highly visible furrow on the seafloor most likely caused by an otter board of a demersal fish trawl, which ran in a straight line exactly matching the trajectory of a recent line identified from penguin tracks. We noted high abundances of benthic scavengers associated with fisheries-related bottom disturbance. While our data demonstrate the acute way-finding capabilities of benthic foraging yellow-eyed penguins, they also highlight how hidden cascading effects of coastal fisheries may alter behaviour and potentially even population dynamics of marine predators, an often overlooked fact in the examination of fisheries’ impacts.     

Seasonal change in foraging areas and dive depths of breeding king penguins at Heard Island

The seasonal variation in the foraging behaviour of king penguins (Aptenodytes patagonicus) was studied at Heard Island (53°05′S, 73°30′E) during 1992/1993. On seven occasions throughout the breeding cycle, time-depth-light recorders were deployed on breeding adults to record the dive activities and foraging. Foraging locations changed with season: in autumn and spring 1992, adults foraged between 48–52°S and 74–78°E, about 370 km NNE of Heard Island close to the Polar Front. Two penguins tracked in winter travelled 2220 km east of Heard Island (95°E) along the northern ice limit, and 1220 km south of Heard Island to approximately 65°S, respectively. In spring (October), the penguins again foraged further north than during winter. The foraging area utilised in October overlapped the area where the penguins foraged in March/April. The penguins' diving behaviour also varied seasonally: the modal depth of deep dives (>50 m) increased from about 100 m in February to 220 m in October. Mean dive depths increased from 70 ± 52 m in March 1992 to 160 ± 68 m in August 1992. Penguins dived deep (>50 m) only during daylight hours (16 h in February, 9 h in July). Mean dive durations ranged from 2.9 ± 1.1 min in March 1992 to 5.1 ± 1.2 min in August 1992. Associated with changes in foraging location and dive behaviour was a change in diet composition: during summer the penguins ingested mainly myctophid fish (>90%) while in winter the most important diet item was squid. Accepted: 19 October 1998     

Energetic cost of foraging in free-diving emperor penguins.

Hypothesizing that emperor penguins (Aptenodytes forsteri) would have higher daily energy expenditures when foraging for their food than when being hand-fed and that the increased expenditure could represent their foraging cost, we measured field metabolic rates (FMR; using doubly labeled water) over 4-d periods when 10 penguins either foraged under sea ice or were not allowed to dive but were fed fish by hand. Surprisingly, penguins did not have higher rates of energy expenditure when they dove and captured their own food than when they did not forage but were given food. Analysis of time-activity and energy budgets indicated that FMR was about 1.7 x BMR (basal metabolic rate) during the 12 h d(-1) that penguins were lying on sea ice. During the remaining 12 h d(-1), which we termed their "foraging period" of the day, the birds were alert and active (standing, preening, walking, and either free diving or being hand-fed), and their FMR was about 4.1 x BMR. This is the lowest cost of foraging estimated to date among the eight penguin species studied. The calculated aerobic diving limit (ADL(C)), determined with the foraging period metabolic rate of 4.1 x BMR and known O(2) stores, was only 2.6 min, which is far less than the 6-min ADL previously measured with postdive lactate analyses in emperors diving under similar conditions. This indicates that calculating ADL(C) from an at-sea or foraging-period metabolic rate in penguins is not appropriate. The relatively low foraging cost for emperor penguins contributes to their relatively low total daily FMR (2.9 x BMR). The allometric relationship for FMR in eight penguin species, including the smallest and largest living representatives, is kJ d(-1)=1,185 kg(0.705).     

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.     

Women from Venus,men from Mars: inter‐sex foraging differences in the imperial cormorant Phalacrocorax atriceps a colonial seabird

Colonial seabirds are central place foragers and likely to be subject to substantial competition for resources. Mechanisms proposed for reducing intra‐specific competition include differential inter‐sex area use mediated by adult choice. We used GPS loggers and dive recorders to study area use and dive depth in a total of 27 male and 26 female imperial cormorants Phalacrocorax atriceps breeding at a colony of some 6500 birds at Punta Leon, Chubut, Argentina during 2004 and 2005. Although time spent travelling and distances between the colony and foraging sites were similar for both sexes, males and females travelled away from their colony using routes virtually perpendicular to each other so that their foraging areas were distinctly different; females hunted close to the coast while males foraged offshore in deeper water. Consideration of foraging efficiency underwater, defined as the duration spent on the bottom divided by the dive cycle duration, showed that females were more efficient at depths < 40 m while males more efficient at depths > 40 m. We suggest that the substantial sexual dimorphism in this species may be responsible for the different depth‐linked foraging efficiencies and that selection for appropriate depths could lead to differential habitat use and putative differences in prey selection.     

Southern rockhopper penguin Eudyptes chrysocome chrysocome foraging at Possession Island

The foraging ecology of rockhopper penguins was studied at Possession Island, southern Indian Ocean, by counting the number of birds departing from and arriving at colonies over the course of the day and by equipping three birds with time/depth loggers, one of which was recovered having recorded a total of 12 days foraging activity. Both the counts and the results from the diving behaviour showed that the birds foraged exclusively diurnally. Maximum dive depth was 66 m although most time was spent between 10 and 25 m, depths that did not accord with the published distribution of their principal prey as detected by nets and acoustics. Received: 29 March 1996/Accepted: 10 June 1996     

Evolutionary and ecological aspects of some Antarctic and sub-Antarctic penguin distributions

Penguins probably originated in the core of Gondwanaland when South America, Africa, and Antarctica were just beginning to separate. As the continents drifted apart, the division filled with what became the southern ocean. One of the remaining land masses moved south and was caught at the pole by the Earth's rotation. It became incrusted with ice and is now known as East Antarctica. Linking it to South America was a series of submerged mountain ranges that formed a necklace of islands. The northern portion of the necklace, called the Scotia Arc, is now the "fertile crescent" of the Southern Ocean. The greatest numbers and biomass of penguins are found here as well as that of krill, the primary prey species of most penguins, and many other marine predators. Today penguins are found throughout the sub-Antarctic islands and around the entire Antarctic continent. Using satellite transmitters and time-depth recorders, while taking advantage of the parental dedication of breeding birds, numerous investigators have described foraging habits of several species of penguins. The information obtained is labor intensive and costly so that studies are restricted to certain species, areas and seasons. Here I review the patterns evident among six of the most abundant and completely studied of the penguins. The variation in behavior is considerable from those species that seldom dive deeper than 20 m in search of prey to those that will dive to depths >500 m to catch mesopelagic fish and squid. Foraging trips from breeding colonies vary among species and with the season. Often the birds travel no more than 30 km and at other times the trips may exceed 600 km. Sub-Antarctic species often reach more productive waters near or within the Antarctic Polar Front zone, where the mixing of Antarctic and sub-Antarctic waters provide rich resources for their prey. Antarctic species usually remain close to shore, along the continental slope, or near the sea ice edge. Less is known about penguins during the pelagic phase between breeding cycles. What we do know is surprising in regard to their dispersal, which ranges from hundreds to thousands of kilometers from the breeding colonies.     

Satellite tracking of male rockhopper penguins Eudyptes chrysocome during the incubation period at the Falkland Islands

The foraging patterns of ten male rockhopper penguins Eudyptes chrysocome from the Falkland Islands were recorded during the incubation period by using satellite telemetry. Irrespective of study site and year, two different foraging areas could be identified. Three foraging trips were directed towards the slope of the Patagonian Shelf c. 140 km to the northeast of the breeding colony and these trips had a duration of 11–15 days. The seven other rockhopper penguins travelled c. 400 km towards the edge of the Falkland Islands' waters; all these foraging trips followed an anti-clockwise direction and lasted 16–27 days. The calculated mean daily travelling speed, based on the time spent underwater and the distance covered between two positions, was significantly higher in birds travelling to the edge of the Falkland Islands' waters compared with those foraging at the shelf slope (4.4±1.6 km/h vs. 3.4±2.0 km/h, respectively). The consistent foraging patterns exhibited during the long trips may be linked to the Falklands current, allowing the penguins to reach remote areas while reducing their energy expenditure. Potential interactions between commercial fisheries and hydrocarbon exploration are discussed.     

Human-induced behaviour in Adélie penguins Pygoscelis adeliae

Summary Adélie penguins Pygoscelis adeliae appear to be little perturbed by man. We examined the incidence of nest desertion and duration of foraging trip in Adélie penguins when manipulated and fitted with devices of differing sizes. Birds with ca. 1 cm clipped from their tail feathers stayed at sea 50% longer than unmarked controls. The length of foraging trip and incidence of nest desertion increased with increasing device volume. Penguins fitted with devices did not reduce foraging trip length to that of unpackaged birds for at least 19 days. The susceptibility of Adélie penguins to disturbance should be carefully considered when activity patterns are being studied.     

Is food availability limiting African Penguins Spheniscus demersus at Boulders? A comparison of foraging effort at mainland and island colonies

The African Penguin Spheniscus demersus (Vulnerable) formed three new colonies during the 1980s, two on the South African mainland (Stony Point and Boulders) and one on Robben Island. One of the mainland colonies, at Boulders, Simon's Town, is in a suburban area, resulting in conflict with humans. Growth of the Boulders colony was initially rapid, largely through immigration, but has since slowed, possibly as a result of density‐dependent effects either on land (where there has been active management to limit the spread of the colony) or at sea. We test the latter hypothesis by comparing the foraging effort of Penguins feeding small chicks at island and mainland sites, and relate this to the foraging area available to birds. Three‐dimensional foraging paths of African Penguins were reconstructed using GPS and time–depth loggers. There were no intercolony differences in the rate at which birds dived during the day (33 dives/h), in diving depths (mean 17 m, max. 69 m) or in travelling speeds. The maximum speed recorded was 2.85 m/s, with birds travelling faster when commuting (average 1.18 m/s) than when foraging (0.93 m/s) or resting at sea (0.66 m/s during the day, 0.41 m/s at night). There were strong correlations between foraging trip duration, foraging range and total distance travelled. Foraging effort was correlated with chick age at Robben Island, but not at Boulders. Contrary to Ashmole's hypothesis, birds from Boulders (c. 1000 pairs) travelled further (46–53 km) and foraged for longer (13.2 h) than did birds from Robben Island (c. 7000 pairs) and Dassen Island (c. 21 000 pairs) (33 km, 10.3 h). The mean foraging range also differed significantly between mainland (18–20 km) and island colonies (9 km). The area available to central‐place‐foraging seabirds breeding on the mainland is typically less than that for seabirds breeding on islands, but the greater foraging range of Boulders birds results in an absolute foraging area roughly twice that of island colonies, and the area per pair is an order of magnitude greater for the relatively small Boulders colony. Ashmole's hypothesis assumes relatively uniform prey availability among colonies, but our results suggest this does not apply in this case. The greater foraging effort of Boulders birds probably reflects reduced prey availability in False Bay, and thus the recent slowing in growth at the colony may be the result of differential immigration rather than management actions to limit the spatial growth of the colony.     

Seasonal and annual variation in the diving behaviour of Hutton's shearwater (Puffinus huttoni)

ABSTRACT

With the development and implementation of tracking technology, we are now able to monitor the foraging behaviour of seabirds while at sea. Time-Depth Recorders (TDRs) were fitted to Hutton's shearwaters (Puffinus huttoni), an endangered endemic New Zealand species, to measure how diving behaviour varies over the breeding cycle. Hutton's shearwaters (～350?g) dive up to 339 times per day (average 68.8) at depths to 35?m (average 5.6?m), and for periods up to 60?s (average 19.2?s). Incubating birds dived deeper than birds feeding chicks, and a significant difference in diving depth and dive duration were detected at different times of the day. Neither dive frequency nor dive duration differed significantly between years, but there was some annual variation in dive depths. The temporal variation we observed in the diving behaviour of Hutton's shearwaters suggests they are likely to exploit different types of pelagic prey at different stages in their breeding cycle. With on-going changes in the marine environment, monitoring changes in feeding behaviour using TDRs may provide a way to assess environmental change and improve the conservation of this species.     

Overnight foraging trips by chick‐rearing Nazca Boobies Sula granti and the risk of attack by predatory fish

Most tropical booby species complete breeding foraging trips within daylight hours, thus avoiding nights at sea. Nazca Boobies Sula granti are unusual in this respect, frequently spending one or more nights away from the nest. We used GPS dataloggers, time‐depth recorders, and changes in body weight to characterize foraging trips and to evaluate potential influences on the decisions of 64 adult Nazca Boobies to spend a night at sea, or to return to their chicks on Isla Española, Galápagos, in daylight hours. The tagged birds foraged east of Isla Española, undertaking both single‐day (2–15 h, 67% of trips) and overnight trips (28 h–7.2 days, 33%), and executing 1–19 foraging plunge‐dives per single‐day trip. Birds might forage longer if they are in nutritional stress when they depart, but body weight at departure was not correlated with trip length. Birds might be expected to return from longer trips with more prey for young, but they returned from single‐day and overnight trips with similar body weights, consistent with previous indications that Nazca Boobies forage until accumulating a target value of prey weight. Birds with a lower dive frequency during the first 5 h of a trip were more likely to spend the night at sea, suggesting that they might choose to spend the night at sea if prey capture success was low. At night, birds almost never dived and spent most of their time resting on the water’s surface (11.8–12.1 h, > 99% of the time between civil sunset and civil dawn). Thus, the night is an unproductive time spent among subsurface predators under low illumination. The birds’ webbed feet provided evidence of this risk: 24% of birds were missing > 25% of their foot tissue, probably due to attacks by predatory fish, and the amount of foot tissue lost increased with age, consistent with a cumulative risk across the lifespan. In contrast, other tropical boobies (Blue‐footed Sula nebouxii and Brown Boobies Sula leucogaster), which do not spend the night on the water, showed no such damage. These results suggest that chick‐rearing Nazca Boobies accept nocturnal predation risk on occasions of low prey encounter during a foraging trip’s first day.