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     

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.     

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.     

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.     

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.     

Diving characteristics of southern rockhopper penguins (Eudyptes c. chrysocome) in the southwest Atlantic

The diving behaviour of southern rockhopper penguins (Eudyptes c. chrysocome) was studied at two breeding sites in the Southwest Atlantic: the Falkland Islands and Staten Island, Argentina. Incubating and brooding birds were equipped with time-depth recorders to monitor their foraging activities. Rockhopper penguins from Staten Island started their breeding season about 3 weeks earlier than their conspecifics from the Falkland Islands. The foraging area used by incubating males from the Falkland Islands comprised about 150,000 km² to the northeast of the breeding site and was characterised by shelf and slope waters, whereas the foraging area of incubating males from Staten Island comprised 350,000 km² of oceanic waters to the southeast of the breeding site. A number of dive parameters were measured and compared between the four study groups: Incubating males and brooding females from the Falkland Islands, and incubating males and females from Staten Island. In all study groups, dive depth correlated positively to light intensity, dive duration and vertical velocity. However, significant differences between various diving parameters of the study groups were noted, not only in terms of diving performance, but also as regards diving efficiency (DE). A principal component analysis (PCA) on 16 variables revealed that 75% of the variance could be explained by only two principal components: diving pattern (PC1) and diving effort (PC2). PC1 indicated that the birds from Staten Island, both males and females, dived deeper, covered a greater vertical distance per hour and had higher ascent rates, but spent less time underwater and at the bottom of a dive, and had a lower DE than conspecifics from the Falkland Islands. PC2, which included the percentage of foraging dives, the number of dives per hour, dive duration, bottom time and descent rate, differed significantly between incubating males from the Falkland Islands and the other three groups, which were all very similar. Overall, the diving behaviour was notably similar to that of conspecifics from the Indian and Pacific Oceans. The implications of the results in terms of intra-specific adaptations as well as potential threats from human activities are discussed.     

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.     

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.     

Variation in foraging effort by lactating Antarctic fur seals: response to simulated increased foraging costs

Seasonally breeding predators, which are limited in the time available for provisioning young at a central location, and by the fasting abilities of the young, are likely to maximize energy delivery to the young by maximizing the rate of energy delivery averaged over the whole period of investment. Reduction in food availability or increased foraging costs will alter the optimal behavior of individuals. This study examined the behavioral adaptations of a diving predator, the Antarctic fur seal, to increased foraging costs during lactation. One group of mothers (n=5, treatment) was fitted with additional drag to increase the cost of transport in comparison with a control group (n=8). At the scales of the individual dives, the treatment group made more shorter, shallower (< 30 m) dives. Compensation for slower swimming speeds was achieved by diving at a steeper angle. Overall, diving behavior conformed to several specific theoretical predictions but there were also departures from theory, particularly concerning swimming speed during diving. Diving behavior appears to be adjusted to maximize the proportion of time spent at the bottom of dives. At the scale of diving bouts, no difference was observed between the treatment and control groups in terms of the frequency and duration of bouts and there was also no difference between the two groups in terms of the proportion of time spent diving. At the scale of complete foraging cycles, time taken to return to the pup was significantly longer in the treatment group but there was no difference in the rate of delivery of energy (measured from pup growth rate) to the pups in each group. Since mothers in the treatment group did not use significantly more body reserves, we conclude that behavioral adjustments at the scale of individual dives allowed mothers in the treatment group to compensate for the additional foraging costs. Pup growth rate appears to be less sensitive to the foraging conditions experienced by mothers than foraging trip duration. Received: 14 June 1996 / Accepted after revision: 16 November 1996     

Satellite tracking of Humboldt penguins (Spheniscus humboldti) in northern Chile

During the El Niño of 1982/1983, the Humboldt penguin population diminished dramatically in the whole distributional area of the species. Recovery of the population was slow since 1983 and it has been suggested that large numbers of Humboldt penguins die at sea, entangled in nets, or starve to death, even during non-“El Niño” years. We were able to determine for the first time, how Humboldt penguins on Pan de Azúcar Island (26°S; 72°W) utilize their marine habitat and where their feeding areas lie. For this purpose we employed two streamlined Argos satellite transmitters during the 1994/1995 and 1995/1996 breeding seasons, respectively. Mean travelling speed of Humboldt penguins during foraging trips was 0.94?m s^?1 and 50% of bird positions were located within 5?km of the island (90% within 35?km). Total area covered by Humboldt penguins foraging from Pan de Azúcar Island was 12?255?km². Satellite transmitters also recorded dive duration; penguins spent on average 7.8 to 9?h diving per foraging day but showed no preferences for particular feeding areas. Mean daily dive durations (4-d mean) recorded during the 1994/1995 breeding season were positively correlated between birds. Significant correlation between dive duration and sea surface temperature anomalies and negative correlation between dive duration and fishery landings at nearby Caldera harbour indicate that the 1994/1995 increase in foraging effort was a response to deteriorating prey availability. Sea surface temperatures during the 1995/1996 breeding season were colder than average, and we observed no trends in bird diving activities.     

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.     

Flexible foraging strategies of gentoo penguins <Emphasis Type="Italic">Pygoscelis papua</Emphasis> over 5 years in the South Shetland Islands,Antarctica

Gentoo penguins Pygoscelis papua show considerable plasticity in their diet, diving, and foraging behaviors among colonies; we expected that they might exhibit similar variability over time, at a single site, since flexible foraging habits would provide a buffer against changes in prey availability. We examined interannual changes in the foraging strategies and diet of gentoo penguins in the South Shetland Islands, Antarctica, over 5 years with variable prey abundance. Antarctic krill Euphausia superba was the primary diet item, and fish the secondary, though the importance of these items varied among years. Diving behavior also varied over time: different dive depth distributions were observed among years. Nonetheless, chick-rearing success remained relatively constant, indicating that gentoo penguins were able to maintain chick provisioning by altering their foraging strategy among years. Variable abundance of krill in the region did not have observable impacts on the diet, foraging behaviors or chick-rearing success of gentoo penguins. We suggest that foraging plasticity may be one reason that gentoo penguin populations have remained stable in the region, while their congeners (P. antarctica and P. adeliae) with less flexible foraging strategies have declined.     

Depth utilisation by breeding Adélie penguins,Pygoscelis adeliae,at Esperanza Bay,Antarctica

Miniature depth gauges were attached in December 1987 and January 1988 to Adélie penguins,Pygoscelis adeliae, breeding at Esperanza on the Antarctic Peninsula. Results from 34 birds showed that foraging penguins with eggs and with brooded and crèching chicks spent mean periods away from the nest of 96, 36 and 21 h, respectively, during which time means of 29.0 h (30%), 11.2 h (31%) and 2.7 h (13%), respectively, were spent under water at depths > 5 m. Time under water was positively correlated with time absent from nest. Maximum depth reached was 170 m but overall birds spent most time at shallower depths. Birds foraging for brooded chicks dived deeper than birds foraging for crèching chicks. Stomach-pumping indicated that the principal prey caught at this time was krill,Euphausia superba. Mean mass changes of adults during single foraging trips indicated that krill were caught at a mean rate of 7.2 g min^–1 spent under water.     

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.     

Crossing the frontier: vertical transit rates of deep diving cormorants reveal depth zone of neutral buoyancy

Knowing the depth zone of neutral buoyancy of divers is important because buoyancy can determine how animals manage their energy budget. In this study, we estimate the depth zone of neutral buoyancy of free-ranging cormorants for the first time, using time-depth recorders. We discovered that vertical ascent rates of 12 Crozet and 15 Kerguelen diving blue-eyed shags (respectively Phalacrocorax melanogenis and P. verrucosus) slowed down considerably at the 50–60 m depth zone. We suggest this was due to birds trying to reach the surface from that point upwards using reduced locomotor activity because the force of buoyancy becomes greater than the force of gravity at that depth. The results show a shift of this depth zone in relation to maximum targeted dive depth, suggesting cormorants may control buoyancy through respiratory air volume adjustment. Interestingly, 60 m is close to the maximum depth zone reached by these two species during dives lasting 4 min, their estimated behavioural aerobic dive limit. This suggests that the decision to swim deeper has a direct consequence on the energy budget, with time spent recovering at the surface (time thus lost to foraging) strongly increasing relative to the preceding time of submergence. Resources found in deeper waters must be of sufficient quantity or quality to justify crossing the frontier of physical neutral buoyancy.     

Going deep: common murres dive into frigid water for aggregated,persistent and slow-moving capelin

Owing to the necessity of delivering food to offspring at colonies, breeding seabirds are highly constrained in their foraging options. To minimize constraints imposed by central-place foraging and to optimize foraging behavior, many species exhibit flexible foraging tactics. Here we document the behavioral flexibility of pursuit-diving common murres Uria aalge when foraging on female capelin Mallotus villosus in the northwest Atlantic. Quite unexpectedly, being visual foragers, we found that common murres dived throughout the day and night. Twenty-one percent of recorded dives (n = 272 of 1,308 dives) were deep (≥50 m; maximum depth = 152 m, maximum duration = 212 s), bringing murres into sub-0°C water in the Cold Intermediate Layer (CIL; 40–180 m) of the Labrador Current. Deep dives occurred almost exclusively during the day when murres would have encountered spatially predictable aggregations of capelin between 100 and 150 m in the water column. Temperatures within the CIL shaped trophic interactions and involved trade-offs for both predators and prey. Sub-0°C temperatures limit a fish’s ability to escape from endothermic predators by reducing burst/escape speeds and also lengthening the time needed to recover from burst-type activity. Thus, while deep diving may be energetically costly, it likely increases certainty of prey capture. Decreased murre foraging efficiency at night (indicated by an increase in the number of dives per bout) reflects both lower light conditions and changing prey behavior, as capelin migrate to warmer surface waters at night where their potential to escape from avian predators could increase.     

The influence of subsurface thermal structure on the diving behavior of northern fur seals (<Emphasis Type="Italic">Callorhinus ursinus</Emphasis>) during the breeding season

In the heterogeneous marine environment, predators can increase foraging success by targeting physical oceanographic features, which often aggregate prey. For northern fur seals (Callorhinus ursinus), two prevalent oceanographic features characterize foraging areas during summer in the Bering Sea: a stable thermocline and a subsurface “cold pool”. The objective of this study was to examine the influence of these features on foraging behavior by equipping fur seals from St. Paul Island (Alaska, USA) with time-depth recorders that also measured water temperature. Foraging bout variables (e.g., mean dive depth and percent time diving in a bout) were compared with respect to subsurface thermal characteristics (thermocline presence and strength and cold pool presence). Over 74% of bouts occurred in association with strong thermoclines (temperature change > 5°C). Few differences were found for dive behavior in relation to the presence of a thermocline and the cold pool, but for epipelagic bouts, a strong thermocline resulted in increased bottom times, number of dive wiggles, and percent time diving when compared to moderate thermoclines. There was also a positive relationship between mean dive depth and thermocline depth. The combination of increasing foraging effort in areas with strong thermoclines and diving to depths closely related to the thermocline indicates this feature is important foraging habitat for northern fur seals and may act to concentrate prey and increase foraging success. By recognizing the environmental features northern fur seals use to find prey, managers will be better equipped to identify and protect foraging habitat that is important to northern fur seals, and possibly other marine predators in the Bering Sea.     

Reproduction and distribution of the invasive European fanworm Sabella spallanzanii (Polychaeta: Sabellidae) in Port Phillip Bay, Victoria, Australia

 The European fanworm Sabella spallanzanii (Gmelin, 1791) was recently introduced to Port Phillip Bay and is now a conspicuous component of most benthic communities. Reproduction of the worm was investigated in a population at Queenscliff over a 2 yr period (October 1995 to October 1997) using gonadal histology. The worms are dioecious (sex ratio 1:1, n=250), and attained sexual maturity at ∼50 mm body length. Reproductive periodicity followed a distinct annual cycle, and spawning proceeded through an extended autumn/winter period. Spawning was broadly synchronous between sexes, and coincided with falling seawater temperatures and shorter day-lengths. The females were highly fecund, and >50 000 eggs were probably shed from large females (>300 mm body length) during the annual spawning period. Breeding cycles of S. spallanzanii in Port Phillip Bay are ∼6 mo out of phase with endemic populations located at similar latitudes in the northern hemisphere. The spread of S. spallanzanii within Port Phillip Bay has been monitored by divers on an annual basis since 1994. The most recent dive survey (1998) indicates that S. spallanzanii has extended its range through out the entire 2000 km² embayment, and has invaded most subtidal habitats. Quantitative estimates of S. spallanzanii abundances were highest on pier pylons (12.5 individuals m⁻², 0.5 to 7 m depths). On sediments, estimates were highest at shallow sites (0.3 m⁻², 7 m depth), but numbers declined significantly with depth (0.1 m⁻², 17 to 22 m depth). Mean worm lengths and biomass were, by contrast, significantly higher at intermediate depths (12 to 17 m) than in shallower (7 m) or deeper (22 m) locations. S. spallanzanii demonstrates a clear preference for growth in sheltered, nutrient-enriched waters, so it may not spread from Port Phillip Bay into the adjacent oceanic waters of Bass Strait; however, in view of S. spallanzanii's current high abundance, fecundity and extended spawning periodicity, there is a high risk of future range expansions, mediated by shipping, into other temperate-water ports. Received: 17 November 1998 / Accepted: 6 January 2000     

Temporal changes in activity budgets of chick-rearing Adélie penguins

Changes in sea-ice conditions can affect locomotion on land, diving behavior, and corresponding foraging success of penguins. In this study, locomotion on land and diving behavior were compared between early and late stages of the guard phase with different sea-ice conditions using miniaturized time-depth-acceleration data loggers for Adélie penguins Pygoscelis adeliae from 18 December 2001 to 11 January 2002 in Dumont d’Urville, Adélie Land (66.7°S, 140.0°E), Antarctica. Differences were found between early and late stages in the ratio of walking vs. tobogganing, proportion of time spent diving, diving depth as well as in the rate of parental tissue accumulation. In contrast, trip duration, distance traveled on land, and meal delivery rate to chicks did not differ between the stages. This study suggests that physical changes in sea-ice during the penguins’ chick-rearing period may affect certain on-land and/or at-sea behaviors which, in turn, may affect how resources are allocated to self-maintenance or chick-provisioning.     

Foraging behaviour of sympatric Antarctic and subantarctic fur seals: does their contrasting duration of lactation make a difference?

The duration of periods spent ashore versus foraging at sea, diving behaviour, and diet of lactating female Antarctic (Arctocephalus gazella, AFS) and subantarctic (A. tropicalis, SFS) fur seals were compared at Iles Crozet, where both species coexist. The large disparity in lactation duration (SFS: 10 months, AFS: 4 months), even under local sympatry, has led to the expectation that AFS should exhibit higher foraging effort or efficiency per unit time than SFS to allow them to wean their pups in a shorter period of time. Previous evidence, however, has not supported these expectations. In this study, the distribution of foraging trip durations revealed two types of trips: overnight (OFT, <1 day) and long (LFT, >1 day), in common with other results from Macquarie Island. However, diving behaviour differed significantly between foraging trip types, with greater diving effort in OFTs than in LFTs, and diving behaviour differed between fur seal species. OFTs were more frequent in SFS (48%) than in AFS (28%). SFS performed longer LFTs and maternal attendances than AFS, but spent a smaller proportion of their foraging cycle at sea (66.2 vs. 77.5%, respectively). SFS dove deeper and for longer periods than AFS, in both OFTs and LFTs, although indices of diving effort were similar between species. Diel variation in diving behaviour was lower among SFS, which foraged at greater depths during most of the night time available than AFS. The diving behaviour of AFS suggests they followed the nychthemeral migration of their prey more closely. Concomitant with the differences in diving behaviour, AFS and SFS fed on the same prey species, but in different proportions of three myctophid fish (Gymnoscopelus fraseri, G. piabilis, and G. nicholsi) that represented most of their diet. The estimated size of the most important fish consumed did not vary significantly between fur seal species, suggesting that the difference in dive depth was mostly a result of changes in the relative abundance of these myctophids. The energy content of these fish at Iles Crozet may thus influence the amount and quality of milk delivered to pups of each fur seal species. These results contrast with those found at other sites where both species coexist, and revealed a scale of variation in foraging behaviour which did not affect their effort while at sea, but that may be a major determinant of foraging efficiency and, consequently, maternal investment.