Sex‐specific prey partitioning in breeding piscivorous birds examined via a novel,noninvasive approach

Piscivorous birds frequently display sex‐specific differences in their hunting and feeding behavior, which lead to diverging impacts on prey populations. Cormorants (Phalacrocoracidae), for example, were previously studied to examine dietary differences between the sexes and males were found to consume larger fish in coastal areas during autumn and winter. However, information on prey partitioning during breeding and generally on sex‐specific foraging in inland waters is missing. Here, we assess sex‐specific prey choice of Great Cormorants (Phalacrocorax carbo) during two subsequent breeding seasons in the Central European Alpine foreland, an area characterized by numerous stagnant and flowing waters in close proximity to each other. We developed a unique, noninvasive approach and applied it to regurgitated pellets: molecular cormorant sexing combined with molecular fish identification and fish‐length regression analysis performed on prey hard parts. Altogether, 364 pellets delivered information on both, bird sex, and consumed prey. The sexes differed significantly in their overall prey composition, even though Perca fluviatilis, Rutilus rutilus, and Coregonus spp. represented the main food source for both. Albeit prey composition did not indicate the use of different water bodies by the sexes, male diet was characterized by higher prey diversity within a pellet and the consumption of larger fish. The current findings show that female and male cormorants to some extent target the available prey spectrum at different levels. Finally, the comprehensive and noninvasive approach has great potential for application in studies of other piscivorous bird species.     

Diet analysis in piscivorous birds: What can the addition of molecular tools offer?

In trophic studies on piscivorous birds, it is vital to know which kind of dietary sample provides the information of interest and how the prey can be identified reliably and efficiently. Often, noninvasively obtained dietary samples such as regurgitated pellets, feces, and regurgitated fish samples are the preferred source of information. Fish prey has usually been identified via morphological analysis of undigested hard parts, but molecular approaches are being increasingly used for this purpose. What remains unknown, however, is which dietary sample type is best suited for molecular diet analysis and how the molecular results compare to those obtained by morphological analysis. Pellets, feces, and regurgitated fish samples of Great Cormorants (Phalacrocorax carbo sinensis) were examined for prey using both morphological hard part analysis and molecular prey identification. The sample types and methods were compared regarding number of species detected (overall and per sample) as well as the prey species composition and its variability among individual samples. Via molecular analysis, significantly higher numbers of prey species were detected in pellets, feces, and fish samples. Of the three sample types, pellets contained the most comprehensive trophic information and could be obtained with the lowest sampling effort. Contrastingly, dietary information obtained from feces was least informative and most variable. For all sample types, the molecular approach outperformed morphological hard part identification regarding the detectable prey spectrum and prey species composition. We recommend the use of pellets in combination with molecular prey identification to study the diet of piscivorous birds.     

OBSERVATIONS ON THE NESTING SITES AND NESTING BEHAVIOUR OF THE KITTLITZ'S SANDPLOVER CHARADRIUS PECUARIUS

Jackson, S. 1984. Predation by Pied Kingfishers and Whitebreasted Cormorants on fish in the Kosi estuary system. Ostrich 55:113-132.

Identification of otoliths from the regurgitated pellets of Pied Kingfishers Ceryle rudis and Whitebreasted Cormorants Phalacrocorax carbo from the Kosi estuary system provides information on the relative proportions of fish species in the diets of the birds. This information can be related to the feeding habits, distribution and abundance of their prey. It is also an indication of the feeding range of the birds. There is little overlap between both the size classes and the species of fish taken by the two predators. This is because of the difference in size and fishing techniques of C. rudis and P. carbo, and of differences in their feeding ranges. Competition for food between the two populations of birds studied is minimized by these differences.     

Otolith chemistry of prey fish consumed by a fish predator: does digestion hinder Russian doll techniques?

The effect of digestion by a predatory fish (largemouth bass Micropterus salmoides) on stable isotopic (δ¹³C and δ¹⁸O) and trace elemental (Sr:Ca and Ba:Ca) compositions of prey fish (bluegill Lepomis macrochirus) otoliths was investigated in a laboratory experiment. Trace element and stable‐isotopic signatures of L. macrochirus otoliths were not significantly altered for up to 16 h after L. macrochirus were consumed by M. salmoides. Prey fish otoliths recovered from predator digesta can retain environmental stable isotopic and trace elemental signatures, suggesting that determination of environmental history for prey fishes by stable‐isotope and trace‐element analysis of otoliths recovered from stomachs of piscivorous fishes will be feasible.     

Do flocks of great cormorants and goosanders avoid spatial overlap in foraging habitat during the non-breeding season?

Species distribution, ecology, and behaviour are shaped, amongst other things, by interspecific, antagonistic interactions, and this phenomenon is particularly noticeable among predators. We studied the spatial co-distribution of two top piscivorous bird species foraging on inland waters outside breeding season. We considered the hypothesis that goosanders, Mergus merganser, and great cormorants, Phalacrocorax carbo, avoid foraging in close proximity to each other. Data collected on five river-reservoir systems in the Western Carpathians (Poland and Slovakia) during two periods (2014–2015 and 2015–2016) showed that goosander numbers reduced significantly and their foraging areas changed when large flocks of cormorants arrived and began foraging. We also found that inter-flock distances were greatest between flocks of goosanders and cormorants, suggesting that the former species avoided the waters occupied by the latter. Distribution of flocks of both species was additionally determined by the location of foraging place in river-reservoir system, weather, and presence of other piscivorous birds (e.g. grebes) and raptors (e.g. eagles). Together with the results of research in adjacent Bohemia, this study suggests that competition between cormorants and goosanders may arise when bodies of water suitable for piscivorous foraging are scattered and limited in number, as in the mountainous areas of Central Europe.     

Foraging behaviour and prey size of the painted stork

The foraging behaviour of painted stork Mycteria leucocephala was studied during 2004–2006 at 14 different sites in the Delhi region, India. Observations were recorded on 131 individuals, including 29 juvenile birds using a video camera. Recordings were also made at the nesting colony in Delhi zoo to study the prey sizes regurgitated to nestlings. The results confirm that the painted stork is a tactile forager and exclusively piscivorous. Foraging group size ranged from 1 to 18 individuals. Per 5 min foot stirring rates in the vegetated habitats were significantly higher than in non-vegetated habitats. The attempt rate and feeding rate in the breeding season were significantly higher than that in the non-breeding season. Prey sizes taken in the breeding season were significantly smaller than those taken in the non-breeding season. About 80% fish fed to the chicks were smaller than 10 cm. Young chicks were offered smaller prey compared with older chicks. The variations in foraging parameters are discussed in relation to habitats and their conservation in the Delhi region.     

How important are pelagic preys for the kelp gull during chick-rearing at the South Shetland Islands?

An analysis of pellets regurgitated indicated adult kelp gulls (Larus dominicanus) on the South Shetland Islands consumed predominantly intertidal prey, whereas previous studies at Antarctic Peninsula sites have reported kelp gulls consuming predominantly pelagic species. The pellets collected at Nelson Island during the chick-rearing period indicated that the limpet Nacella concinna was their most frequent prey, followed by carrion, gammariids, snails and krill. Fish were scarcely represented. Also, regurgitated stomach contents of chicks showed that limpets and carrion were the most frequent food items, accounting for 70% of the mass. However, gammariids were particularly important by number. Significant differences were observed in the overall comparison of the diet as reflected by the two sampling methods. In general, the importance of pelagic prey was negligible when compared to intertidal or scavenged prey. Our results differ greatly from those reported for the Antarctic Peninsula, where chicks were almost exclusively fed with the pelagic fish Pleuragramma antarcticum. These differences could be related to the abundance of pelagic resources in southernmost latitudes, and/or to the presence of more extensive intertidal foraging areas at the South Shetland Islands. Received: 7 October 1996 / Accepted: 14 July 1997     

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.     

How equally sized piscivorous birds and fish sharing common food resources may reduce possible feeding interactions between them?

The diets of sympatric predators may overlap, especially when their body sizes are similar and foraging area is relatively small. It may be also supposed that some differences in their foraging strategies may counteract competitive interactions among them, and therefore be of advantage to these species. To reveal such phenomena the composition of food of cormorant and adult pikeperch was studied in the Dobczyce Reservoir (S Poland) from June to November 2002. The main prey species were the same and the range of prey size was similar for both piscivores. Despite these similarities, the potential for dietary overlap was strongly reduced due to two differences in their foraging patterns: (1) different preferred prey species (cormorants foraged mainly roach, whereas pikeperch selected juvenile percids); (2) different size of simultaneously selected prey (in summer, cormorants selected larger prey, while in autumn larger prey was selected by pikeperch). These differences may be explained by some general features of birds and fishes, which determine the costs to the individual of capturing prey. The observed selection of different prey species and sizes may be also important for the co-occurrence of other piscivorous birds and fishes sharing common food resources.     

Commensal association of the common kingfisher with foraging Eurasian otters

Positive interactions between birds and mammals are a fascinating aspect of animal behaviour. Feeding associations may consist of local enhancement or facilitation, and in the latter case, of commensalism or mutualism depending on the benefits received by the facilitator. We report here on a previously undescribed feeding association between piscivorous birds and Eurasian otters Lutra lutra. In Spain, common kingfisher Alcedo atthis and grey heron Ardea cinerea were observed closely following foraging otters and benefited from feeding opportunities provided by these. Behavioural observations of otters in central Spain (28.4 hr; 19 days) revealed that an association with kingfishers occurred in 33% of otter foraging events (n = 92). Simultaneous observations in northern Spain (14.2 hr; 16 days) showed an association between otters and kingfishers or grey herons in 41.6% and 11.7% of otter foraging events (n = 77), respectively. The association probability between kingfishers and otters increased significantly when otters foraged closer to the shore and on small fish rather than other prey (crayfish or large fish). Birds fed on prey remain left by the feeding otters, on small fish captured by otters when these were satiated and playing, or on prey displaced by otters. Our observations are consistent with facilitation and commensalism: piscivorous birds gained feeding opportunities provided by the otters, with no apparent costs or benefits to the latter. Similar feeding associations have been described between other species of otters and piscivorous birds (kingfishers, herons, egrets, storks) in Asia, South America and Southern Africa, but had not yet been described in Europe. The occurrence of piscivorous bird–otter associations in different species and regions suggests that this commensalism may be often overlooked but widespread. We have shown that the association can be frequent and is context‐dependent, with benefits for associating birds depending on otters´ behaviour and targeted prey.     

From mass of body elements to fish biomass: a direct method to quantify food intake of fish eating birds

The winter diet of great crested grebe (Podiceps cristatus) was studied in the major lakes of Insubria region, N Italy, in 2000–2003 by analysing stomach and oesophagus contents of birds found dead. Winter diet of cormorant (Phalacrocorax carbo) was studied using pellet analysis. Diet was expressed in terms of numerical abundance, frequency and biomass of prey. A detailed methodology is described of how to prepare a reference collection of otoliths, pharyngeal bones and chewing pads (key-elements), essential for species-specific identification of fish remains in stomachs and pellets, and to estimate ingested fish biomass. Correlations between mass (mg) of key-elements and mass of corresponding fish were significant, and exponential regression models were used to calculate biomass of each single prey item directly from the mass of its key-elements. Advantages of this method are compared with the more traditional two step approach in which size of elements is used to estimate fish length, followed by calculating fresh mass from fish length. Cyprinids formed the largest part of the diet for both predators. Great crested grebes selected small prey, feeding mainly on bleak (Alburnus alburnus alborella), while diet composition of cormorants varied significantly in relation to prey availability. Handling editor: K. Martens     

Foraging patterns and kleptoparasitism among three sympatric cormorants (Phalacrocorax spp.) from the Delhi region, North India

Cormorants, described as ‘foot-propelled pursuit divers’, constitute an important component of aquatic food webs and exhibit unique foraging behaviour patterns, which can be properly understood through a comparative study. Since, after a foraging dive they surface to ingest the prey, the intensity of kleptoparasitic attacks on the surface can have a major impact upon the net energetic gain for each individual. Inspite of the fact that cormorants and their habitats are severely threatened in India, their foraging behaviour has not been adequately studied. Such considerations prompted us to undertake field studies on three sympatric cormorants (Phalacrocorax spp.) at 60 different sites in the Delhi region of North India, during 2004–2007. By means of video photography, some key foraging parameters including group size, prey size and patterns of kleptoparasitic attacks were quantified. Along a loose body size gradient, we observed significant differences among the three species with respect to not only their preference for wetland size but also prey size. The frequency of a kleptoparasitic attacks depended upon the group size and foraging behaviour of each species. It was observed that several foraging bouts were abruptly terminated due to human disturbances, mostly at sites lying outside the protected areas. This observation points towards the need to conserve small waterbodies in the countryside, currently threatened by pollution and urbanization, for the benefit of a variety of waterbirds including cormorants.     

Die Nahrung von Kormoranen (Phalacrocorax carbo sinensis) in Bayern

Zusammenfassung Die wachsende Zahl von Kormoranen (Phalacrocorax carbo sinensis) in Bayern führte in den letzten Jahren zu erheblichen Konflikten zwischen Fischerei und Naturschutz. Deshalb wurde die Zusammensetzung der Kormorannahrung an verschiedenen Gewässertypen untersucht: Zwei große Voralpenseen (Chiemsee, Ammersee), drei künstlich angelegte Gewässer (Altmühlsee, Baggersee Ochsenanger bzw. Breitengüßbacher Baggersee), zwei freifließende bzw. staugeregelte große Fließgewässer (Donau, Inn) und ein kleineres Fließgewässer (Alz). Insgesamt wurden 4697 Speiballen untersucht. Diese enthielten die Hartteile von 9587 Fischindividuen.Von 50 in den Untersuchungsgewässern vorkommenden Fischarten wurden 24 in den Speiballen der Kormorane angetroffen. Allerdings machten nur 6–10 Arten 89–95% aller gefressenen Fischindividuen aus, wobei in den einzelnen Gewässern lediglich 2–3 Arten mit 57–85% die Hauptbeute darstellten. Am häufigsten wurden Weißfische (Fam. Cyprinidae) in den Speiballen nachgewiesen. An den großen Voralpenseen bildete die Renke (Coregonus spec.) zeitweise einen weiteren wichtigen Nahrungsbestandteil. An Ammersee, Altmühlsee, Donau und Inn wurden häufig auch Barsche (Perca fluviatilis) gefressen, an der Donau auch Kaulbarsche (Gymnocephalus cernua) bzw. Schrätzer (G. schraetzer). Dagegen spielten an der schneller fließenden Alz Äschen (Thymallus thymallus) und Salmoniden (Fam. Salmonidae) eine größere Rolle. Außer an der Alz dominierten an allen Untersuchungsgewässern häufige Schwarmfische. Die Nahrung wies an den Voralpenseen und an der Donau die größte Diversität auf.Die Nahrungszusammensetzung variierte häufig von Monat zu Monat. Diese Fluktuationen weisen auf Verhaltensweisen bzw. horizontale und vertikale Wanderbewegungen der einzelnen Fischarten hin. Häufig zeigen sie auch einen Wechsel der von den Vögeln befischten Habitate auf.Meist wurden Fische mit Körperlängen von 9–28 cm erbeutet. An der schneller fließenden Alz wurden im Mittel größere Fischen gefangen als in allen übrigen Gewässern. Am Ammersee, dem einzigen Gewässer das ganzjährig untersucht werden konnte, wurden in den Sommermonaten kleinere Fische erbeutet als im Winter. Nur wenige Jungfische wurden gefressen. So wiesen lediglich 32 Speiballen (1%) mehr als 20 Fische auf.

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The food of Cormorants (Phalacrocorax carbo sinensis) in Bavaria

Summary In Bavaria, an increase in Cormorant (Phalacrocorax carbo sinensis) numbers has lead to increasing conflict between fishermen and conservationists. The diet of cormorants was assessed by examining the contents of oral pellets from seven study sites, thought to be representative of Bavarian waters. These were two large pre-alpine lakes (Chiemsee and Ammersee), a reservoir (Altmühlsee), two gravel pits (Ochsenanger and Breitengüßbacher Baggersee), two uncontrolled or controlled large rivers (Danube and Inn), and a smaller river (Alz). The sample of pellets comprised 2944 collected from these sites in the years 1991–1994 and 1758 from an earlier study (winter 1990/91). Altogether. 9587 individual prey items were found. Of 50 fish species inhabiting the study waters, 24 were found in the pellets of the cormorants, but only 6–10 species accounted for 89–95% by numbers. Moreover. within each site 2 or 3 species predominated by number (57–85%).Most fish remains in pellets were from cyprinids (Cyprinidae) but at times on pre-alpine lakes whitefish (Coregonus spec.) were consumed in numbers. At the lakes Ammersee and Altmühlsee, as well as on the rivers Danube and Inn, cormorants also fed commonly on Perch (Perca fluviatilis), and along the Danube also on Ruffe (Gymnocephalus cernua) and Striped Ruffe (G. schraetzer), respectively. Only at the River Alz, with its faster flow, were Grayling (Thymallus thymallus) and salmonids (Salmonidae) important as cormorant prey. In general, common shoaling fish species dominated the diet of the birds, an exception being the River Alz again. The diet showed highest diversity at the pre-alpine lakes and at the River Danube.There was some seasonal variation in the prey species composition. This was probably caused by fish behaviour and fish movements, which perhaps influenced their vulnerability to cormorant predation. Some variation also suggested birds switched between feeding sites. Cormorants used a wide range of the available prey sizes. Mostly, fish of 9–28 cm in length were taken. At the River Alz, the fish sizes consumed were larger than at all other sites. At Lake Ammersee, the only water studied year-round, larger prey were taken in winter than in summer. In general, few fry were eaten, as only 32 pellets (1%) contained more than 20 individual fish.

     

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.     

Small-scale spatial variation in the elemental composition of otoliths of Stegastes nigricans (Pomacentridae) in French Polynesia

Solution-based inductively coupled plasma-mass spectrometry (ICP-MS) was used to determine if Stegastes nigricans collected from 15 sites in French Polynesia could be distinguished by the trace element composition of their otoliths. A total of 293 adults were collected by spearing and their otoliths were analysed. We found that elemental signatures differed significantly among sites within and between the islands of Tahiti and Moorea (p<0.001), primarily due to variation in concentrations of the elements Ba, Ca, Li, Mg, Mn, Na, Sr and Y. The otoliths of fish collected within Papeete Harbour in Tahiti had distinctive elemental signatures characterised by relatively high concentrations of Mn. Otoliths of these fish could be distinguished from others that were collected only a small distance (200 m) from the harbour. This is the first time that differences in chemical composition of otoliths have been reported at such small spatial scales and this trait may prove useful for the studies of connectivity of populations at within reef scales.     

The diet of Herring Gulls (Larus argentatus) during winter and early spring on the lower Great Lakes

In the Great Lakes, the Herring Gull (Larus argentatus) is a prominent member of the aquatic bird community, and has been used to monitor spatial and temporal trends in contaminant levels. To understand more fully contaminant loading outside the breeding season, we analysed the contents of 1298 freshly regurgitated pellets and 179 fresh faeces, collected in March and early April 1978–83, and between late December and late February 1990–91, from the vicinity of breeding colonies in Lakes Ontario and Erie, the Niagara River, Detroit River, and south-eastern parts of Lake Huron. Most adult Herring Gulls from the Great Lakes population winter in these areas, but there is no published account of their food habits other than during the breeding season. Most pellets from colonies close to large urban centres contained remains of garbage, as well as various fish species. Small mammals, notably Deer Mice (Peromyscus maniculatus) dominated the early spring diet at Lake Huron colonies near agricultural areas. At all other sites fish predominated in pellets and faeces, but garbage items were also recorded regularly. The species of fish consumed varied regionally, probably reflecting local availability. In Lake Ontario, Rainbow Smelt (Osmerus mordax) and Alewife (Alosa pseudoharengus) occurred most frequently in samples, whereas Freshwater Drum (Aplodinotus grunniens) was the main fish prey in Lake Erie and the Detroit River. Dietary differences were apparent between years, within seasons, and amongst areas. While these may have reflected some real differences in food availability, interpretation of these results was confounded by various biases inherent in the sampling of pellets and faeces to determine diet in such an opportunistic species. Therefore, it would be unwise to draw rigid conclusions as to regional or seasonal differences in the diets of piscivorous birds, based upon analyses of diet from only a small sample of sites or years. Herring Gulls appear to feed mainly on fish and garbage in winter and early spring on the lower Great Lakes (much as during the breeding season), but any locally abundant food source is probably exploited opportunistically.     

Diet composition and food consumption of the grey heron (Ardea cinerea) from breeding colonies in northern Poland

The diet of the grey heron (Ardea cinerea) was investigated in three colonies in northern Poland (Mosty, Kiersity and Kty Rybackie) varying in size, habitat type and composition of feeding areas used by birds. The diet was assessed during the breeding season on the base of pellets and regurgitated food. Pellets from all studied colonies consisted mainly of mammal hair (almost 100% of samples), bone remains (20–24%) and invertebrate remains (26–51%, mainly water beetle, Dytiscus marginalis). Bones and hair of water vole (Arvicola terrestris) and Microtus voles were the most common remains of mammals recorded in pellets. Remains of other animal taxa were found sporadically. In the colonies at Mosty and Kiersity, fish composed more than 95% of collected regurgitated prey items. Food consumption of herons from the biggest colony at Kty Rybackie was estimated to exceed 100 t per season. Herons from Mosty consumed ca 46–52 t, from Kiersity 30–38 t of food per season. The impact of herons on fish communities was much lower than other losses due to predation (herons from Kty Rybackie caught only 6% of fishermen bycatch, birds from Mosty took 10–14% of round gobies eaten by cormorants).     

Flexible foraging techniques in breeding Cormorants Phalacrocorax carbo and Shags Phalacrocorax aristotelis: benthic or pelagic feeding?

A total of 8772 dive durations were recorded during 117 diving bouts in five Cormorants Phalacrocorax carbo and five Shags Phalacrocorax aristotelis breeding at the Chausey Islands, France. Diet of the birds was assessed by analysis of 526 pellets containing 13,016 otoliths. Radio-tracking data indicated that Cormorants fed exclusively on pelagic fish during social fishing (5% of the trips) and executed 11% pelagic and 60% benthic dives during the remaining 95% of the trips. In Shags, 44% of all trips were pelagic, and the remaining 56% included 9% pelagic and 67% benthic dives. The proportions of benthic to pelagic dives varied widely between dive sequences of single birds and between individuals and sexes in both species. The prey spectrum of the Cormorants contained both pelagic (29%) and benthic fish (67%) and confirmed considerable flexibility in foraging. In Shags, birds may adjust their diving patterns to accommodate the behaviour of their main prey, sandeels Ammodytidae (87% of all prey). We propose that the wetability of plumage may explain this flexibility.     

Sex and individual differences in foraging behavior of Japanese cormorants in years of different prey availability

Japanese cormorants, Phalacrocorax capillatus, are sexually dimorphic seabirds with males that are heavier and that dive deeper than females. Sex differences in prey composition and foraging behavior of those rearing chicks at Teuri Island, Hokkaido, were examined by collecting food-loads from parents in 1992–1998 and by radio-tracking ten birds each in 1997 and 1998 when prey availability was different. Males fed more on benthic and epibenthic fishes (82% mass) than did females (34%) while females fed more on epipelagic and coastal fishes (53%) than did males (18%). Males made longer dives (53 s) at feeding sites closer to the island (7 km) than females (39 s, 13 km) in 1997. In 1998 when the availability of epipelagic fish seemed to be higher, there were no sex differences in dive duration and distance to the feeding sites (35 s and 9 km for males, 36 s and 10 km for females). This sex difference in foraging behavior with a poor availability of epipelagic fish suggests that high diving ability possibly enables males to feed on demersal fish. Birds specializing in coastal shallow waters around the island made long dives; hence they were probably foraging in bottom layers. Those foraging in deeper shelf waters made short dives and they were thought to forage in surface layers. Electronic Publication     

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.