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.     

Luxury in freshwater and stress at sea? The foraging of the Common Tern Sterna hirundo

We compared the foraging strategies of Common Terns Sterna hirundo in freshwater (Lake Jeziorsko, Brzeg, Poland) and marine environments (Minsener Oldeoog, German Wadden Sea). Body mass changes, nest relief and duration and number of feeding trips per day were studied by automatically weighing the adults, using electronic balances under the nests. At the freshwater site, adults were lighter both before and after feeding and gained less mass during a trip. in the Wadden Sea, single feeding trips lasted longer than at the freshwater site and the terns made fewer trips per day. To achieve the same mass gain per day as in birds in freshwater, trips at sea had to be longer and food intake per trip was higher. The daily duration of absence for feeding and the daily mass gain were about the same in both areas. The limnetic feeders finished foraging earlier in the evening than the terns foraging at sea. These differences are consistent with the hypothesis that limnetic prey availability was consistent, whereas the tides limited the availability of marine prey. In consequence, foraging over freshwater presents several advantages, such as higher colony attendance, better mate coordination and better parental care.     

Behaviour and parental care of Skylark Alauda arvensis chicks

I studied parental care and fledgling dispersion of Skylarks Alauda arvensis during the last 4 days of the nestling period and the first 4 days after the nestlings left the nest but before they were able to fly. The study compared parental care in three different crop types: spring barley, grass and set-aside. Both parents made more provisioning trips and delivered more food to fledglings than to nestlings. Fledglings received fewer items per trip than did nestlings. Feeding distances did not differ between nestling and post-Hedging periods for any crop type. This suggests that fledging was associated with a change in parental foraging strategies. Parental care differed between broods from different crop types. During the last 4 days of the nestling period, the feeding frequencies were 3.4 trips per young per hour in spring barley fields, 5.8 in grass and 7.3 in set-aside. The mean distances to the feeding area were 233 m in spring barley fields, 155 m in grass and 120 m in set-aside. The load size of provisioning trips was significantly higher in set-aside than in spring barley and grass. During the first 4 days after fledging, the feeding frequencies were 2.2 successful trips per young per hour in spring barley fields, 4.1 in grass and 5.1 in set-aside. The feeding distances were 210 m in spring barley fields, 162 m in grass and 120 m in set-aside. Load size of provisioning trips was significantly higher in set-aside than in spring barley and grass. The mean dispersion of fledglings was significantly greater in fields of spring barley compared with grass fields and set-aside.     

Foraging trip decisions by the streaked shearwater Calonectris leucomelas depend on both parental and chick state

Parents of albatross and shearwater species employ a dual foraging strategy, feeding their chicks quickly in repeated short trips and then restoring their own fuel reserves during longer trips. A decline in parental body condition is believed to trigger longer trips, but chick body condition and age may also play a role. To investigate these factors in the little-studied streaked shearwater Calonectris leucomelas, we monitored the nest attendance of 17 pairs on Mikura Island in 2005 using an automated identification system. We also monitored body mass changes and meal masses of 5 of the 17 pairs using an automated weighing system. Although the birds did not show a clear dual foraging pattern, trip duration varied widely from 1 to 15 days. On average, the birds fed chicks 67.6 g during nighttime meals at 2.74-day intervals. Since meal mass did not depend on trip duration, feeding efficiency (meal mass delivered per unit trip duration) decreased as trip duration increased. Parents accumulated more energy reserves when they took longer trips. Parents appeared likely to initiate longer trips when their body condition declined or chick body condition recovered.     

Variability of mouth width in European eel,Anguilla anguilla,in relation to varying feeding conditions in three Dutch lakes

Synopsis Mouth widths of eels in three Dutch lakes were compared relative to feeding conditions over two years. Average mouth width varied by season and year and per lake. Of two groups of feeding specialists distinguished in each population chironomid feeders showed a smaller mouth width compared to fish feeders of the same body length. Proportions of these specialists differed between years, lakes, and seasons and changed in response to changing feeding conditions. The shift to fish feeding occurred when chironomid biomass decreased and young fish recruited. The shift was greatest in the year when chironomids collapsed completely and in the lakes with high eel densities. Mature males were narrow-headed and left the lakes at sizes between 30 and 40 cm. Larger eels were always broad-headed females. Implications for foraging efficiency and population structure are discussed.     

Between-lake differences in the diet and provisioning behaviour of Black-throated Divers Gavia arctica breeding in Scotland

Surveys at 23 Black-throated Diver breeding lakes in Scotland showed large between-lake differences in the species and size range of potential prey. The study lakes were classified into four types according to the main size-taxa prey classes present. Type 1 lakes lacked small fish, Type 4 lacked salmonids < 300 mm and Types 2 and 3 both had all four of the main prey classes. Diver diet for 30 families at the study lakes was quantified from 7943 prey items seen fed to chicks, and 153 items seen eaten by adults, during 662 h of observations. At all lake types adults mostly ate salmonids of 120–240 mm in length, especially Brown Trout Salmo trutta , and European Eels Anguilla anguilla up to c.  350 mm. The prey eaten by chicks ranged widely, from c.  0.03 g to 300 g, with important implications for feeding behaviour. Young chicks (days 1–8) rejected items greater than 70 mm long. The diet of young chicks consisted mostly of small fish, either Three-spined Sticklebacks Gasterosteus aculeatus (Type 2 lakes) or Minnow Phoxinus phoxinus (Type 3 and Type 4 lakes). At other lakes (Type 1) it consisted mostly of mayfly larvae (Ephemeroptera); invertebrates were the only abundant potential small prey in these lakes. At all lakes the chick diet became more similar to that of adults as chicks grew. Adults provisioning chicks used two foraging strategies. In 'excursion foraging', adults hunted away from the chicks and carried prey back for them; in 'attendance foraging', the chicks accompanied the foraging adult(s). Excursion foraging was used mostly before day 4 and was strongly and positively correlated with the abundance of fish < 80 mm in length. This behaviour is probably adaptive, but is probably tenable only where small fish are relatively abundant. Young chicks fed mainly on small fish had higher survival rates than those fed on invertebrates.     

Does the foraging strategy of adult Short‐tailed Shearwaters cause obesity in their chicks?

The chick provisioning behaviour of Short-tailed Shearwaters Puffinus tenuirostris breeding at the northern edge of their distribution on Montague Island, New South Wales, was examined in February and March 1997. The duration of individual foraging trips of parents, weight changes of adults and chicks, and meal sizes delivered to chicks were determined. It was found that individual parents mixed a long foraging trip to Antarctic waters (14.4±2.0 days) with one to three short foraging trips (1.36±0.7 days, mode=1 day). Adults gained body mass on long trips and lost weight on short trips. The size of meals fed to the chicks was significantly greater after a long trip (161±21 g) than after a short trip (135±28 g), although short trips increased the overall chick feeding frequency. The variable number of short trips made by adult Short-tailed Shearwaters and the relationship between short trips and adult body condition were consistent with current life-history theory: adults do not sacrifice their own body condition to increase food delivered to their chicks.
Modelling revealed that this dual foraging strategy inevitable leads to chicks enduring long intervals between meals. These long intervals may have led to the evolution of an over-feeding strategy by parents and the nestling obesity reported in this shearwater. The durations of the long trips from Montague Island were significantly greater than those for Short-tailed Shearwaters breeding at the centre of their distribution in Tasmania, although there was no significant difference in the length of short trips. A commitment to feed regularly in Antarctic waters may explain why the breeding distribution of this species does not extend much further north.     

The foraging decisions of a central place foraging seabird in response to fluctuations in local prey conditions

During reproduction, seabirds need to balance the demands of self- and offspring-provisioning within the constraints imposed by central place foraging. To assess behavioral adjustments and tolerances to these constraints, we studied the feeding tactics and reproductive success of common murres (also known as common guillemots) Uria aalge , at their largest and most offshore colony (Funk Island) where parents travel long distances to deliver a single capelin Mallotus villosus to their chicks. We assessed changes in the distance murres traveled from the colony, their proximate foraging locations and prey size choice during two successive years in which capelin exhibited an order of magnitude decrease in density and a shift from aggregated (2004) to dispersed (2005) distributions. When capelin availability was low (2005), parental murres increased their maximum foraging distances by 35% (60 to 81 km) and delivered significantly larger capelin to chicks, as predicted by central place foraging theory. Murres preferred large (>140 mm) relative to small capelin (100–140 mm) in both years, but unexpectedly this preference increased as the relative density of large capelin decreased. We conclude that single prey-loading murres target larger capelin during long foraging trips as parents are 'forced' to select the best prey for their offspring. Low fledgling masses suggest also that increased foraging time when capelin is scarce may come at a cost to the chicks (i.e. fewer meals per day). Murres at this colony may be functioning near physiological limits above which further or sustained adjustments in foraging effort could compromise the life-time reproductive success of this long-lived seabird.     

Foraging behaviour and feeding locations of Rock Shags Phalacrocorax magellanicus from a colony in Patagonia, Argentina

During 1996 and 1997, foraging Rock Shags Phalacrocorax magellanicus were studied at Punta Loma, Argentina using radio-transmitters deployed on ten adult shags during the chick-rearing period. Rock Shags undertook 2.6 ± 0.6 sd trips per day. The mean duration of a feeding trip was 2.6 ± 0.7 hours. A bird spent 36% of daylight hours away from the colony on feeding trips, diving for 92% of the foraging trip, and made a mean of 106 dives per foraging trip. Foraging trip duration was strongly correlated with the total number of dives made in one foraging trip. Rock Shags fed mainly in water less than 10m deep with a gravelly sand bottom and within 5 km of shore. Mean foraging range was 3.8 ± 2.6 km and 2.6 ± 2.3 km for 1996 and 1997, respectively. These results suggest a high foraging effort (diving time per foraging trip) for Rock Shags, presumably associated with poor food conditions close to the colony. Comparison is made with other Phalacrocorax species.     

The development of regular seasonal habitat shifts in a landlocked Arctic charr, Salvelinus alpinus L., population

Parts of the Arctic charr population of the subarctic Lake Visjön in north-west Sweden migrate upstream during the spring to two small, recently eutrophied and very productive lakes. Large repeat migrants arrive first, followed by young first-time migrants. Charr in the small lakes grow more rapidly than those resident in L. Visjön. In early September mature fish leave the lakes, followed by immature fish later in September and in October. Overwintering and spawning takes place in L. Visjön. Migratory females attain maturity at age 4 years and resident females at age 6 years. The migrant fish return annually until they are 5–6 years old. This limit may be due to reduced relative growth benefits of the habitat shift for larger individuals. The rapid development of these regular habitat shifts could be explained by an internally fixed exploratory behaviour in these Arctic charr that makes the detection and utilization of distant feeding resources possible. Migrants will possess a considerably higher fitness, if survival rates for migratory and resident fish are equal.     

BREEDING BIOLOGY OF THE BEARDED VULTURE IN SOUTHERN AFRICA,PART III: THE POST-NESTLING DEPENDENCE PERIOD

Brown, C. J. 1990. Breeding biology of the Bearded Vulture in southern Africa, Part III: The post-nestling dependence period. Ostrich 6l: 43–49.

The post-nestling dependence period of the Bearded Vulture Gypaetus barbatus in southern Africa begins with the first flight of the young bird at 126 ± 2 days after latching (November-January) and ends during the pre-laying nod or the parent birds' next breeding attempt (April-June), a nod of about five months. For the first two weeks after first flit young bid remaine6 within about 200m of the nest, moving up to 800 m by the third week. By a month out of the nest young birds spent about 40% of the day in flight, moved up to 3 km from the nest, began bone-dropping and interacting with young birds from neighbouring nests. At six weeks they began to accompany their parents for part of some of their foraging trips, but returned to the nest alone, and by eight weeks they completed foraging forays with parents Pasting up to 3 h. At 2–3 months out of the nest young birds covered an area of about 42 km², excluding the foraging trips with parents, by 3–4 months, 78km² and 4–6 months, 168 km². Parent birds delivered food for at least five months after the young bird's first flight. Young birds left their natal areas of their own accord, usually during the first month of their parents' next breeding attempt.     

Foraging patterns of breeding Dolphin Gulls Larus scoresbii at Punta Tombo, Argentina

The Dolphin Gull Larus scoresbii is a little-known, rare species endemic to southern South America. Knowledge of its feeding ecology is essential for development of management and conservation strategies. To obtain information on their use of food resources and on the frequency and duration of their foraging trips, we followed seven individuals by radiotelemetry during the breeding season of 2002 at the Punta Tombo Reserve, Argentina. In 99% of mapped locations ( n  = 4069) the Gulls were within 2.4 km of their colony. Dolphin Gulls foraged mostly at the colonies of three other species, feeding on Magellanic Penguin Spheniscus magellanicus and Imperial Cormorant Phalacrocorax atriceps regurgitates and on Southern Sea Lion Otaria flavescens faeces. The sea lion colony was the most frequently visited foraging area, accounting for 64% of the total ( n  = 260 trips). A similar trend was observed during both the incubation (60%, n  = 38 trips) and the chick (64%, n  = 40 trips) stages. The mean number of trips per day was 3.8 (range = 3–5), with a mean duration of 123 min (sd = ±27). Trip duration differed between foraging areas, being significantly longer at the sea lion than at the penguin and cormorant colonies. The small foraging range given by its particular feeding strategy suggests that Dolphin Gulls, in contrast to most other seabirds that range over large expanses of ocean in search of food, may be effectively conserved within protected areas during the breeding season.     

Time constraint on food choice in provisioning blue tits, Parus caeruleus: the relationship between feeding rate and prey size

Previous work on food-provisioning behaviour in blue tits suggested that the parents could gather larger prey items only by making longer foraging excursions, for example, by being more selective or by reaching more distant (and less exploited) feeding sites. Here, I show that within-nest, within-day variation in size of prey delivered by the parent could be explained by the time since its last visit. In unmanipulated conditions, size of larvae tended to increase with the time spent away from the nest. A significant positive relationship was more likely at high provisioning rates, suggesting that periods of intense feeding limited the size of prey delivered to the brood. To assess the effect of less intense feeding on prey size, I experimentally increased food availability to the tits. The parents could decide whether to eat the extra food or feed it to the nestlings. In both cases, food supplementation could result in longer time lags between natural feedings. Food-supplemented parents consumed the extra food and fed it to their nestlings, made longer foraging trips and delivered larger natural larvae than controls. In this group, size of larvae was more constant during the observation period and was independent of the time since the parent's last visit. This suggests that, below some value of visit rate, prey size is no longer limited by the duration of the foraging trip. The results support the view that tits continually vary visit rate and prey size. There is some evidence that these adjustments are made by changing food selectivity in response to changes in the state of the brood and of the parents.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Diet and diving behaviour of European Storm Petrels Hydrobates pelagicus in the Mediterranean (ssp. melitensis)

Capsule Unlike Atlantic populations, which feed on krill, Mediterranean populations feed mainly on pelagic fish Gymnammodites cicerellus.

Aims To determine the diet and dive depth of the Mediterranean subspecies of European Storm Petrels Hydrobates pelagicus melitensis.

Methods Analysis of regurgitates of adults arriving at the colony for chick feeding and by determination of dives depth using the capillary tube method.

Results The main prey is Gymnammodites cicerellus, a pelagic fish. Storm Petrels dive for their prey and can reach up to 5 m in depth. They also make short foraging trips just outside the colony where they capture Opossum Shrimps Misydacea.

Conclusions European Storm Petrels in the Mediterranean exploit pelagic fish which are taken by diving. This contrasts with the Atlantic populations which feed mainly on krill. Mediterranean birds also feed on Opossum Shrimps Mysidacea during short foraging trips made at night just outside the colony. Differences in diet between long and short foraging trips may be because adults have to forage for both themselves and their chicks.     

Diets and feeding of Fulmars Fulmarus glacialis during the breeding season:a comparison between St Kilda and Shetland colonies

Diets and nest attendance of Fulmars at Foula, Shetland and St Kilda, Outer Hebrides were examined during the breeding season. At Foula, sandeels formed the bulk of the diet, with fish offal and pelagic zooplankton of minor importance. At St Kilda, 71% of regurgitates consisted of pelagic zooplankton, probably captured at night. Dietary overlap between the two colonies was 14% by species composition. Foraging trips from Foula shortly after chick hatching generally lasted for less than 10 h, while trips from St Kilda late in chick rearing often lasted more than 24 h. The diurnal pattern of feeding suggested mainly nocturnal foraging from St Kilda but both diurnal and nocturnal foraging from Foula.     

Similarities in feeding behaviour between some marine and freshwater fishes in two tropical communities

The feeding behaviour and diets of fishes in two tropical habitats, a marine reef and a freshwater pond, were studied comparatively in Brazil. Similarities were found in the tactics employed to obtain food, the social patterns during foraging, and the general diet, notwithstanding lower-level taxonomic differences between the food items. The feeding behaviours of about one-third of the fish fauna from each community were approximately equivalent. The feeding categories of these fishes are briefly described. The similarities in the feeding modes probably reflect structural and functional properties shared by the two communities. Additional behavioural similarities of fishes in both habitats are presented and the lack of some particular foraging modes in each community is noted. The picture emerged that different, unrelated fish assemblages have the ability to evolve towards a similar behavioural and structural organization in response to comparable situations and constraints. The value of underwater observations and naturalistic studies on tropical freshwater fish assemblages is indicated.     

Spatial and temporal variation in the provisioning behaviour of female rockhopper penguins Eudyptes chrysocome filholi

As seabirds are central place foragers during breeding, their provisioning behaviour and their ability to face variable energy demand from the chicks is expected to vary with environmental conditions. The provisioning behaviour of female rockhopper penguins Eudyptes chrysocome filholi was recorded over the chick‐rearing period at Kerguelen (KER) and Crozet (CRO) archipelagoes (two very distinct marine environments), using time‐depth recorders, or VHF transmitters coupled with an automatic recording station. No influences of the method have been found on the average foraging trip durations. Some previously undescribed short and multiple trips within a day were recorded using the automatic recording system. These multiple trips (6.8 h) were mostly performed with <5 days old chicks, a period during which feeding rates were the highest (1.1 meals per day), at both sites. During the brooding period, both KER and CRO females mainly performed daily trips of increasing duration (2 h longer at CRO) and at decreasing frequency. During the crèche compared to the brooding period, females from KER performed slightly fewer daily trips (0.6 per day) and more (<3 days) overnight trips, while females from CRO performed very few daily trips (0.1 per day) and more overnight trips, some of them being long trips lasting 5 to 29 days, mostly initiated during the transition between the brooding and the crèche periods. The result fit the hypothesis that long trips permit females to restore and/or maintain their body condition at more distant foraging places. It seemed that chick developement during the brooding period and environmental factors during the crèche period conditioned trip duration of females. Due to more long trips at CRO, the female feeding frequency was twice as high at KER than at CRO during the crèche period, while males participated in the feeding duties. Based on differences in female behaviour, we hypothesize that the male's contribution is likely to differ strongly from one site to another, and may buffer the possible decrease in female feeding frequency by feeding the chicks if food is less abundant.     

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

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

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.     

Nest attendance and foraging movements of northern fulmars rearing chicks at Bjørnøya Barents Sea

We studied several aspects of the foraging ecology of fulmars rearing young chicks on Bjørnøya. To determine precisely the duration of foraging trips during the brooding period, we used an automated logging system that recorded the presence of fulmars fitted with transponders. We also tracked, with satellite transmitters, four parent fulmars during the brooding period, and two after the chick had been left alone. When brooding the chick, fulmars appeared to alternate very rapidly on the nest, with foraging trips lasting on average 8?h. This period appeared constraining for the birds since parents lost mass. The growth of chicks was dependent on the ability of the female (and not the male) to do short foraging trips. At this time birds are foraging at an average distance of 60?km from the colony, with birds concentrating on the shelf around Bjørnøya. They did not return from one trip to the next to the same foraging area. As the season progressed and the chicks were left alone on the nest, parents increased the duration and maximum range of foraging trips as well as the distance covered. However, they still perform a succession of relatively short foraging trips to the east of the Bjørnøya shelf but they interspersed these short trips with longer foraging trips. One bird returned twice to the same site along the Norwegian coast 570?km from Bjørnøya, the other foraged at 580?km in the mid-Barents Sea. Average flight speed including time spent on the water was 28?km/h and reached 70?km/h during bouts of more than 1?h when the bird was probably continuously in flight.