Sex‐specific food provisioning in a monomorphic seabird,the common guillemot Uria aalge: nest defence,foraging efficiency or parental effort?

Sexual differences in food provisioning rates of monomorphic seabirds are well known but poorly understood. Here, we address three hypotheses that attempt to explain female-biased food provisioning in common guillemots Uria aalge : (1) males spend more time in nest defence, (2) females have greater foraging efficiency, and (3) males allocate a greater proportion of foraging effort to self-maintenance. We found that males spent no more time with chicks than females but made longer trips and travelled further from the colony. There was extensive overlap between sexes in core foraging areas, indicating that females were not excluding males from feeding opportunities close to the colony. However, as a result of their longer trips, the total foraging areas of males were much greater than those of females. There was no difference between sexes in overall dive rate per hour at sea, in behaviour during individual dives or in a number of other measures of foraging efficiency including the frequency, depth and duration of dives and the dive: pause ratio during the final dive bout of each trip, which was presumably used by both sexes to obtain prey for the chick. These data strongly suggest that sexes did not differ in their ability to locate and capture prey. Yet males made almost twice as many dives per trip as females, suggesting that males made more dives than females for their own benefit. These results support the hypothesis that female-biased food provisioning arose from a difference between sexes in the allocation of foraging effort between parents and offspring, in anticipation of a prolonged period of male-only post-fledging care of the chick, and not from differences in foraging efficiency or time spent in nest defence.     

Foraging behaviour of Razorbills Alca torda during chick-rearing at the largest colony in the Baltic Sea

Capsule: Foraging behaviour in the Razorbill Alca torda during breeding was similar to that found elsewhere, aside from dive shape.

Aims: To investigate the foraging behaviour of Razorbills during the breeding season at the largest colony in the central Baltic Sea.

Methods: A combination of global positioning system (GPS) and time-depth recorder (TDR) devices were used on Razorbills breeding on the island of Stora Karlsö, Baltic Sea, during the chick-rearing period.

Results: Five GPS tracks and nine TDR logs were retrieved from 12 Razorbills, and 7399 dives were analysed. Razorbills foraged south and southwest of the colony. Maximum and mean (±sd) foraging range from the colony was 72.7?km and 13.1?±?13.5?km, respectively. Mean dive depth (15.3?±?2.4?m) and duration (53.1?±?8.5?s) were similar to those of a more southern Baltic Sea Razorbill colony. Dive depth had a bimodal distribution, with 70% of dives deeper than 10?m and 30% shallower than 10?m. There was a clear diel foraging pattern with 89% of dives occurring during daytime and a higher proportion of shallow dives at night. Unexpectedly, dives were primarily U-shaped. The Razorbills spent 31% of their overall time activity budget flying or diving.

Conclusion: Aside from dive shape, foraging behaviour was consistent with that reported at other colonies of Razorbills. Inconsistency in dive shape may be due to a bimodal foraging strategy, local prey behaviour or competition with the Common Guillemot Uria aalge.     

Vertical movement of dolphinfish <Emphasis Type="Italic">Coryphaena hippurus</Emphasis> as recorded by acceleration data-loggers in the northern East China Sea

Environmental changes influence foraging behavior for most animals. Dolphinfish, Coryphaena hippurus, are epipelagic predators and have a cosmopolitan tropical to warm-temperate (>20°C) distribution. We simultaneously obtained the ambient temperature and the foraging behavior (i.e., swimming speed, depth and tailbeat acceleration) of dolphinfish, using an acceleration data-logger in May, September, October, November 2007, June 2008, May and July 2010 for 8 individuals. Although the dolphinfish spent a mean ± standard deviation of 43.4 ± 27.7% of their time at the surface (0–5 m), dive excursions from the surface (DES) were observed in all individuals and maximum DES depths ranged from 50.1 to 95.4 m. DES events resulted dives below the thermocline for these dolphinfish, and there was a significantly positive relationship between the isothermal layer depth (ILD) and DES depth. Our results demonstrate that dolphinfish avoided the rapid thermal change beyond the thermocline, and their prey is most likely found in the upper layers of the thermocline. Gliding behavior during the DES phase was also observed and dolphinfish gradually descended to deeper waters with gliding. The gliding time was longer when the ILD was deeper, and fish tended to dive deeper. We suggest that dolphinfish adopt gliding behavior to search a broader range of depths for prey, while minimizing energy use.     

On a wing and a prayer: the foraging ecology of breeding Cape cormorants

The Cape cormorant Phalacrocorax capensis is unusual among cormorants in using aerial searching to locate patchily distributed pelagic schooling fish. It feeds up to 80 km offshore, often roosts at sea during the day and retains more air in its plumage and is more buoyant than most other cormorants. Despite these adaptations to its pelagic lifestyle, little is known of its foraging ecology. We measured the activity budget and diving ecology of breeding Cape cormorants. All foraging took place during the day, with 3.6 ± 1.3 foraging trips per day, each lasting 85 ± 60 min and comprising 61 ± 53 dives. Dives lasted 21.2 ± 13.9 s (maximum 70 s), attaining an average depth of 10.2 ± 6.7 m (maximum 34 m), but variability in dive depth both within and between foraging trips was considerable. The within-bout variation in dive depth was greater when making shallow dives, suggesting that pelagic prey were targeted mainly when diving to <10 m. Diving ecology and total foraging time were similar to other cormorants, but the time spent flying (122 ± 51 min day⁻¹, 14% of daylight) was greater and more variable than other species. Searching flights lasted up to 1 h, and birds made numerous short flights during foraging bouts, presumably following fast-moving schools of pelagic prey. Compared with the other main seabird predators of pelagic fish in the Benguela region, Cape gannets Morus capensis and African penguins Spheniscus demersus , Cape cormorants made shorter, more frequent foraging trips. Their foraging range while feeding small chicks was 7 ± 6 km (maximum 40 km), similar to penguins (10–20 km), but less than gannets (50–200 km). Successful breeding by large colonies depends on the reliable occurrence of pelagic fish schools within this foraging range.     

When cormorants go fishing: the differing costs of hunting for sedentary and motile prey

Cormorants hunt both benthic (sedentary) and pelagic (motile) prey but it is not known if the energy costs of foraging on these prey differ. We used respirometry to measure the costs of diving in double-crested cormorants (Phalacrocorax auritus) foraging either for sedentary (fish pieces) or motile (juvenile salmon) prey in a deep dive tank. Short dives for sedentary prey were more expensive than dives of similar duration for motile prey (e.g. 20% higher for a 10s dive) whereas the reverse was true for long dives (i.e. long dives for motile prey were more expensive than for sedentary prey). Across dives of all durations, the foraging phase of the dive was more expensive when the birds hunted motile prey, presumably due to pursuit costs. The period of descent in all the dives undertaken appears to have been more expensive when the birds foraged on sedentary prey, probably due to a higher swimming speed during this period.     

Pursuit plunging by northern gannets (Sula bassana) feeding on capelin (Mallotus villosus)

Northern gannets (Sula bassana) are considered to obtain prey usually by rapid, vertical, shallow plunge dives. In order to test this contention and investigate underwater foraging behaviour, we attached two types of data-logging systems to 11 parental northern gannets at Funk Island in the North-Wiest Atlantic. We documented, for the first time to the authors' knowledge, gannets performing long, flat-bottomed, U-shaped dives that involved underwater wing propulsion as well as rapid, shallow, V-shaped dives. The median and maximum dive depths and durations were 4.6 and 22.0 m and 8 and 38 s, respectively. Short, shallow dives were usually V-shaped and dives deeper than 8 m and longer than 10 s were usually U-shaped, including a period at constant depth (varying between 4 and 28s with median 8s). Diving occurred throughout the daylight period and deepest dives were performed during late morning. On the basis of motion sensors in the loggers and food collections from telemetered birds, we concluded that extended, deep dives were directed at deep schools of capelin, a small pelagic fish, and we hypothesized that V-shaped dives were aimed at larger, pelagic fishes and squids. Furthermore, these V-shaped dives allowed the birds to surprise their pelagic prey and this may be critical because the maximum swimming speeds of the prey species may exceed the maximum dive speeds of the birds.     

Sex differences in the diving behaviour of a size-dimorphic capital breeder: the grey seal

Both body size dimorphism and sex differences in the relative costs and benefits associated with acquiring energy for reproduction have been advanced to explain the evolution of sex differences in foraging behaviour. We examined the extent to which these factors influenced sex differences in the diving behaviour of a size-dimorphic, capital breeder, the grey seal, Halichoerus grypus. Using time-depth data loggers, we examined the diving behaviour of 46 male and 49 female grey seals for 7 months before parturition and mating. Males and females showed significantly different seasonal patterns in the characteristics of individual dives and dive effort. Compared with males, females showed significantly higher levels of dive effort immediately following moult and in the 3 months before parturition. Females also had longer dives (5.5 versus 4.9 min) and spent more time at depth (3.4 versus 2.7 min), whereas males dived deeper (57 versus 49 m). Males dived consistently throughout the day, whereas females showed strong diurnal patterns in dive depth, duration and frequency. The diving behaviour and rates of mass gain by females suggested a pattern of foraging consistent with early accumulation of body energy to support pregnancy and the subsequent lactation period during which females fast. Males, on the other hand, showed diving behaviour and rates of mass gain consistent with a more gradual accumulation of energy stores. Our results suggest that sex differences in the seasonal patterns of diving behaviour reflect sex differences in the costs and benefits of stored energy for reproduction rather than the influence of body size dimorphism alone.     

Foraging and breeding performance of Japanese cormorants in relation to prey type

Seabirds are high trophic predators in marine ecosystems and are sensitive to change in food supply and thus seabirds can be used as monitors of the marine environment. In order to study the foraging responses of Japanese cormorants Phalacrocorax filamentosus breeding at Teuri Island, Hokkaido to changes in fish availability, the diet was assessed from the regurgitations of parents and chicks, and diving behavior was measured by using time-depth recorders. Breeding performance (brood size, chick growth, breeding success) was monitored using conventional methods to study their breeding responses. Japanese cormorants changed the diet and foraging behavior over four summers. The birds fed mainly on epipelagic schooling fish when they were available and on demersal fish when pelagic fish availability was low. They tended to dive deeper and longer in a year when they fed mainly on demersal fish than the other years, reflecting the change in the depth distribution of prey fish. Chick growth rate did not differ among years, but fledging success was lower in the years of demersal fish as their meal delivery rate was low. When epipelagic schooling fish were considered scare, parents maintained chick growth by reducing brood size. High variability and unpredictability in pelagic fish abundance are key factors affecting the foraging and breeding performance of Japanese cormorants, which could potentially be used to monitor fish resources.     

Diving and haulout behavior of crabeater seals in the Weddell Sea,Antarctica, during March 1986

Summary Time-depth recorders were used to study the diving and haulout behavior of six crabeater seals in the marginal. ice edge zone of the Weddell Sea during March 1986. Haulout patterns revealed the seals' clear preference for diving during darkness and hauling out onto sea ice during daylight. Seals did not necessarily haul out every day; individual seals hauled out on 80–100% of days during the study period. Four general dive types were identified: 1) traveling dives, 2) foraging dives, 3) crepuscular foraging dives, and 4) exploratory dives. Nearly continual diving occurred for extended periods (about 16 h) nightly, with one individual diving up to 44 h without interruption. Foraging dives occurring during crepuscular periods were deeper than those made during the darkest hours. The authors suggest that the distinct diel pattern of dive timing and depth may be related to possible predator avoidance behavior by the seals' principal prey, Antarctic Krill.     

Sex-specific foraging behaviour in a seabird with reversed sexual dimorphism: the red-footed booby

Most hypotheses attempting to explain the evolution of reversed sexual dimorphism (RSD) assume that size-related differences in foraging ability are of prime importance, but the studies on sex-specific differences in foraging behaviour remain scarce. We compare the foraging behaviour of males and females in a seabird species with a RSD by using several miniaturised activity and telemetry loggers. In red-footed boobies males are 5% smaller and 15% lighter than females, but have a longer tail than females. Both sexes spend similar time on the nest while incubating or brooding. When foraging at sea, males and females spend similar time foraging in oceanic waters, forage in similar areas, spend similar proportion of their foraging trip in flight, and feed on similar prey—flying fishes and flying squids—of similar size. However, compared to males, females range farther during incubation (85 km vs. 50 km), and furthermore feed mostly at the extremity of their foraging trip, whereas males actively forage throughout the trip. Males are much more active than females, landing and diving more often. During the study period, males lost mass, whereas females showed no significant changes. These results indicate that males and females of the red-footed boobies differ in several aspects in their foraging behaviour. Although some differences found in the study may be the direct result of the larger size of females, that is, the slightly higher speeds and deeper depths attained by females, others indicate clearly different foraging strategies between the sexes. The smaller size and longer tail of males confer them a higher agility, and could allow them to occupy a foraging niche different from that of females. The higher foraging effort of males related to its different foraging strategy is probably at the origin of the rapid mass loss of males during the breeding period. These results suggest that foraging differences are probably the reason for the differential breeding investment observed in boobies, and are likely to be involved in the evolution and maintenance of RSD.     

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.     

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.     

Diving behaviour and diet of the blue-eyed shag at South Georgia

Summary This paper describes a concurrent investigation of individual variation in diet, diving patterns and performance of blue-eyed shags Phalacrocorax atriceps breeding at South Georgia. Within one day individual shags exhibited one of three foraging strategies: short diving (4 birds, all dives 120 s) and mixed diving (15 birds, predominantly long but with a few short dives). The mean number of dives per day was significantly higher in shags that only made short dives (mean=172.0, SE=43.2) than birds with a mixed diving strategy (mean=40.5, SE=4.7) and birds that made only long dives (mean=30.8, SE=1.8). Diet was assessed using hard remains recovered from pellets regurgitated by the shags. Small nototheniid fish (c. 10 kJ per item) were by far the commonest prey but most pellets contained additional items. The frequency of pellets with additional items of higher energy value than nototheniid fish (10.c. 900 kJ per item), lower energy value (>1–10 kJ per item) and both higher and lower energy items was strikingly similar to the frequency of shags making long, short and both long and short dives respectively. Predicted aerobic dive limits suggested that during long dives, blue-eyed shags were probably sustained by anaerobic metabolism. Models of prey capture rates demonstrated that for both long and short diving, many items must be caught per dive when birds are feeding on prey at the lower end of the energy range. Predicted capture rates for the commonest recorded prey (small fish) differ markedly between the two diving strategies.     

Deep-diving foraging behaviour of sperm whales (Physeter macrocephalus)

1. Digital tags were used to describe diving and vocal behaviour of sperm whales during 198 complete and partial foraging dives made by 37 individual sperm whales in the Atlantic Ocean, the Gulf of Mexico and the Ligurian Sea. 2. The maximum depth of dive averaged by individual differed across the three regions and was 985 m (SD = 124.3), 644 m (123.4) and 827 m (60.3), respectively. An average dive cycle consisted of a 45 min (6.3) dive with a 9 min (3.0) surface interval, with no significant differences among regions. On average, whales spent greater than 72% of their time in foraging dive cycles. 3. Whales produced regular clicks for 81% (4.1) of a dive and 64% (14.6) of the descent phase. The occurrence of buzz vocalizations (also called 'creaks') as an indicator of the foraging phase of a dive showed no difference in mean prey capture attempts per dive between regions [18 buzzes/dive (7.6)]. Sperm whales descended a mean of 392 m (144) from the start of regular clicking to the first buzz, which supports the hypothesis that regular clicks function as a long-range biosonar. 4. There were no significant differences in the duration of the foraging phase [28 min (6.0)] or percentage of the dive duration in the foraging phase [62% (7.3)] between the three regions, with an overall average proportion of time spent actively encountering prey during dive cycles of 0.53 (0.05). Whales maintained their time in the foraging phase by decreasing transit time for deeper foraging dives. 5. Similarity in foraging behaviour in the three regions and high diving efficiencies suggest that the success of sperm whales as mesopelagic predators is due in part to long-range echolocation of deep prey patches, efficient locomotion and a large aerobic capacity during diving.     

Effects of increased swimming costs on foraging behavior and efficiency of captive Steller sea lions: Evidence for behavioral plasticity in the recovery phase of dives

A significant component of foraging energetics is the cost of locomotion, which for marine animals, is the cost of swimming. Increases in the cost of swimming may have significant impacts on foraging efficiency. Minimizing the cost of swimming can contribute to the optimization of foraging strategies by reducing the energetic cost of foraging. Results of several field studies suggest that an increase in the cost of locomotion may have comparable effects on foraging behavior and efficiency to a decrease in prey availability. We tested the hypothesis that an increased cost of swimming, brought on by increased hydrodynamic drag, has the same effect on dive behavior and efficiency as reduced prey availability under standard locomotion. Experiments were performed using two adult female Steller sea lions at the Alaska SeaLife Center in Seward, AK, using the same animals and general experimental design previously used to test the effects of reduced prey encounter rate on dive behavior and efficiency. Animals were fitted with a drag-inducing harness for half of the 500 simulated foraging dives in order to increase the cost of swimming. Individual dive duration and foraging time were significantly reduced in all cost-increased dives, comparable to the effects of reduced prey encounter rate. However, on a bout-by-bout basis, dive and foraging efficiency were only slightly reduced, which is likely due to an average 50% reduction in post-dive surface recovery duration during cost-increased dives. Increased heat flux across the body surface measured in a parallel study confirmed a significant increase in work during drag-increased dives. These results suggest that sea lions are able to compensate for changes in the cost of foraging and maintain their foraging efficiency by altering their dive strategy over an entire bout of dives when operating well within their aerobic scope.     

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.     

BIOLOGY OF THE BANK CORMORANT,PART 3: FORAGING BEHAVIOUR

Cooper, J. 1985. Biology of the Bank Cormorant, Part 3: Foraging behaviour. Ostrich 56: 86–95.

The Bank Cormorant Phalacrocorax neglectus, endemic to southern Africa, is primarily a solitary inshore forager. Bank Cormorants forage Primarily on the bottom among kelp beds but also may forage over shingle or coarse sand substrates or in midwater. Breeding birds forage up to 9 km from their colony. Little is known of foraging depth but birds may dive as deep as 28 m. Mean dive duration was 44,9 s and ratio of dives to surface rests was 2,18. In most cases prey is swallowed under water, presumably to avoid kleptoparasitism. Bank Cormorants foraged during daylight hours from before sunrise to after sunset. Birds did not forage in exceptionally rough seas. Mean female foraging bout duration (84,3 min) was significantly longer than that of males (68,4 min) in breeding individuals. Breeding males undertook significantly more foraging bouts (3,47 boutdday) than did females (3,02 bouts/day). No significant differences were found between the sexes when total time spent foraging/day by breeding birds was compared. It is not clear why males foraged more often, but for shorter periods, than did females, but the differences may be related to sexual dimorphism, males being larger than females.     

The influence of wave exposure on the foraging activity of marine otter, <Emphasis Type="Italic">Lontra felina</Emphasis> (Molina, 1782) (Carnivora: Mustelidae) in northern Chile

The marine otter Lontra felina has been said to prefer wave-exposed habitats over more protected sites in response to a greater prey abundance in exposed habitat. We examined how the foraging activity of L. felina is affected by the regime of wave exposure and prey availability at Isla Choros, northern Chile. Through focal sampling we recorded time spent by otters in foraging, the duration of dives, and the hunting success on a wave-exposed and a wave-protected site on the island. In addition, we quantified the abundance of prey in both habitats. Marine otters spent more time foraging in the wave-protected site compared with the wave-exposed habitat. Successful dives reached 26.9% in the wave-exposed habitats, and 38.2% in the wave-protected habitat. Foraging dives were 18% shorter in wave-exposed as compared with wave-protected habitat. Numerically, available prey did not differ significantly with habitat. Our results are more consistent with the hypothesis that wave-exposed habitats represent a sub-optimal habitat to foraging marine otters. Marine otters’ use of wave-exposed patches through northern and central Chile coastal areas probably reflects a low availability of suitable protected areas and greater human disturbance of more protected habitat.     

Cheetahs of the deep sea: deep foraging sprints in short-finned pilot whales off Tenerife (Canary Islands)

1. Empirical testing of optimal foraging models for breath-hold divers has been difficult. Here we report data from sound and movement recording DTags placed on 23 short-finned pilot whales off Tenerife to study the foraging strategies used to catch deep-water prey. 2. Day and night foraging dives had a maximum depth and duration of 1018 m and 21 min. Vocal behaviour during dives was consistent with biosonar-based foraging, with long series of echolocation clicks interspersed with buzzes. Similar buzzes have been associated with prey capture attempts in other echolocating species. 3. Foraging dives seemed to adapt to circadian rhythms. Deep dives during the day were deeper, but contained fewer buzzes (median 1), than night-time deep dives (median 5 buzzes). 4. In most deep (540-1019 m) daytime dives with buzzes, a downward directed sprint reaching up to 9 m s(-1) occurred just prior to a buzz and coincided with the deepest point in the dive, suggestive of a chase after escaping prey. 5. A large percentage (10-36%) of the drag-related locomotion cost of these dives (15 min long) is spent in sprinting (19-79 s). This energetic foraging tactic focused on a single or few prey items has not been observed previously in deep-diving mammals but resembles the high-risk/high-gain strategy of some terrestrial hunters such as cheetahs. 6. Deep sprints contrast with the expectation that deep-diving mammals will swim at moderate speeds optimized to reduce oxygen consumption and maximize foraging time at depth. Pilot whales may have developed this tactic to target a deep-water niche formed by large/calorific/fast moving prey such as giant squid.     

Feeding behavior and modeled energetic intake of common loon (Gavia immer) adults and chicks on small lakes with and without fish

We examined the behavior of common loons, Gavia immer (Brünnich), breeding on small, shallow lakes in central Alberta, Canada that were naturally fishless or contained only small-bodied fishes (minnow lake). For both lake types, adults spent >90% of their time on the nesting lake and >50% of their time foraging. Adult loons on fishless lakes dove more frequently, but dives were of shorter duration than loons on lakes with fish. On two intensively studied fishless lakes, adults fed chicks macroinvertebrates, particularly leeches, whereas on a focal minnow lake, fish made up >70% of prey items delivered by adults. Chicks >36 days of age on a minnow lake spent >50% of their time foraging, whereas older chicks on fishless lakes were highly dependent on food provisioning by adults. Models based on observed foraging patterns indicated that prey size was a better predictor of success in meeting energetic requirements than was feeding behavior (e.g., dive rate, dive success). For most models, estimated energetic intake was higher for loons on minnow lakes than on fishless lakes. Our behavioral observations and model results are consistent with surveys in central Alberta that indicate that breeding Common Loons frequently establish territories on small lakes, but that chicks hatched on lakes completely lacking fish rarely fledge and only if sufficient large invertebrates such as leeches are available.