Foraging activity patterns in the goosander (Mergus merganser) and the red-breasted merganser (M. serrator) in relation to patterns of activity in their major prey species

Summary Diel and seasonal foraging activity patterns of goosanders (Mergus merganser L.) and red-breasted mergansers (M. serrator L.) were studied during the breeding season on the Rickleå River in northern Sweden (64°05N). In addition, the locomotor activity patterns of their most important prey species, the river lamprey (Lampetra fluviatilis (L.)) and the three-spined stickleback (Gasterosteus aculeatus L.), were recorded.During spring, as mergansers arrived in the breeding area, both species exhibited a bimodal, diurnal pattern of activity with morning and evening peaks. By June (the spawning period for the river lamprey and the migratory period for the three-spined stickleback), goosander activity had shifted to late in the evening. This new activity peak correlated well with the main activity period in the river lamprey. Red-breasted mergansers remained predominantly diurnal throughout the observation period. Their diurnal activity correlated with the activity of their major prey, the three-spined stickleback.Both merganser species utilize a specialized foraging technique, i.e. probing the river-bottom with the bill, to catch hiding fish more or less at random. This behaviour probably helps them to forage during the relatively bright summer nights and thus prey upon the abundant nocturnal river lamprey. The goosander preys upon the river lamprey to a greater extent than does the red-breasted merganser, thus leading to a temporal segregation in food resource utilization between the two duck species.     

Variation in the morphology of adult Apatemon gracilis Rudolphi, 1819 (Digenea: Strigeidae) reared in different avian hosts

Adult Apatemon gracilis (Rudolphi, 1819) were reared experimentally in three different avian hosts: herring gulls Larus argentatus Gmelin, domestic chicks Gallus gallus (L.) and eider ducklings Somateria mollissima (L.). Comparisons of size, body proportions and fecundity were made between these and specimens obtained from a naturally infected goosander Mergus merganser L. The eider duck proved to be a suitable experimental host, rearing adults of comparable size and fecundity to those from the natural host. The growth and development of the parasite in domestic chicks and herring gulls were significantly reduced. Similar conditions are likely to exist in the natural environment and this should be taken into consideration when choosing experimental hosts for use in the laboratory or when identifying species.     

Living on predictability: modelling the density distribution of efficient foraging seabirds

In areas with regular fishing coastal fleets seabirds may benefit from the predictability of discards from fishing vessels, but it is not clear to what extent birds rely on this predictable resource and whether foraging is synchronized with the diel availability of discards. In this paper we investigate if a typical scavenger species, the yellow‐legged gull Larus michahellis, takes advantage of the temporal and spatial predictability of fish discards in the western Mediterranean Sea. The activity and distribution of the trawling fleet in this area is regulated and very predictable in time and space. We gathered aerial survey data across a relatively large area close to the coast to study the spatial distribution and density of L. michahellis, and modelled the density distribution of the species in relation to several oceanographic, ecological and temporal variables, using two different modelling approaches: MARS (multivariate adaptative regression splines) and GLM (generalized linear models). Our models suggest that the spatial density of trawlers at sea and the time of the day are the best explanatory variables of gull distribution, and that gulls concentrate in areas with vessels mainly during fish discarding time, supporting the hypothesis that gulls optimize time foraging to take advantage of fishery waste predictability. Additional surveys from the main gull roosting sites inshore support this hypothesis, as gulls start leaving to the sea just before fishing is completed and vessels begin discarding fish scraps when back to the harbour. This study represents one of the few examples of applying MARS to density distribution modelling, although its application to marine ecosystems should be conducted with caution because of large areas with real absence data. GLMs have shown to be more adaptable to such kind of data. Our data confirm the importance of fishery waste for L. michahellis, not only as a food resource but also as a major driver of their activity and distribution patterns. The ability of seabirds to predict accurately when a food resource will be available implies that modelling their distribution at sea needs to include such variables, both in spatial and temporal dimensions.     

Foraging rhythm of great crested grebes Podiceps cristatus adjusted to diel variations in the vertical distribution of their prey Osmerus eperlanus in a shallow eutrophic lake in The Netherlands

Summary Great crested grebes Podiceps cristatus only foraged for an hour or two during dawn and again during dusk on Lake IJsselmeer in August-September. During this time of the year the adult birds are in wing moult and temporarily unable to fly. The food of grebes consisted almost exclusively of smelt Osmerus eperlanus, the most numberous pelagic fish. Simultaneous sonar registrations and trawl net fishing showed that smelt moved to the water surface during the twilight periods. During day and night they were concentrated near the bottom. We argue that grebes have the best foraging opportunities during twilight when much of their prey is near the surface, where light intensities allow the fish to be detected and captured. When the smelt are in the upper water layers the distance to the covered to get the prey (i.e. diving time and cost) is also least.     

Modelling Terrestrial and Marine Foraging Habitats in Breeding Audouin's Gulls Larus audouinii: Timing Matters

Although the breeding ecology of Audouin’s gull has been widely studied, its spatial distribution patterns have received little attention. We assessed the foraging movements of 36 GPS-tracked adult Audouin’s gulls breeding at the Ebro Delta (NW Mediterranean), coinciding with the incubation period (May 2011). This also coincided with a trawling moratorium northwards from the colony. We modelled the distribution of the gulls by combining these tracking data with environmental variables (including fishing activities from Vessel Monitoring System, VMS), using Maxent. The modelling range included both marine and terrestrial areas. Models were produced separately for every 2h time interval across the day, and for 2 fishing activity scenarios (workdays vs. weekends), allowing to assess the spatio-temporal distribution patterns of the gulls and the degree of association with fisheries. During workdays, gull distribution at sea fully matched with fishing activities, both trawling (daylight) and purse-seining (nightime). Gulls tended to avoid the area under trawling moratorium, confirming the high influence of fisheries on the distribution patterns of this species. On weekends, gulls made lesser use of the sea and tended to increase the use of rice fields. Overall, Audouin’s gull activity was more intense during dailight hours, although birds also showed nocturnal activity, on both workdays and weekends. Nocturnal patterns at sea were more disperse during the latter, probably because these gulls are able to capture small pelagic fish at night in natural conditions, but tend to congregate around purse-seiners (which would enhance their foraging efficiency) in workdays. These results provide important insight for the management of this species. This is of particular relevance under the current scenario of European fisheries policies, since new regulations are aimed at eliminating discards, and this would likely influence Audouin’s gull populations.     

The diel activity of carabid beetles (Coleoptera) north of the Arctic Circle,with particular reference to Patrobus assimilis Chaud. and Notiophilus aquaticus L.

Summary Diel activity patterns of carabids species caught in pitfall traps in a subalpine birch forest north of the Arctic Circle are described. Patrobus assimilis Chaud. and Notiophilus aquaticus L. were the most common species and were active at all times of the day with peak activity at noon. Seven less common species and the larvae of Notiophilus aquaticus L. showed similar activity patterns. There was a positive correlation between the mean diel variation in temperature and the activity of the different species. Although day to day variations in temperature influenced the level of activity, it had little effect on the daily activity patterns. The adaptive significance of the diurnal activity patterns of carabid beetles in northern areas is discussed in relation to low night temperatures and high predation pressure from birds.     

Identification of Diplostomum spp. in the retina of perch Perca fluviatilis and the lens of roach Rutilus rutilus from the Baltic Sea — an experimental study

Metacercariae of the genus Diplostomum dwelling in the retina of perch Perca fluviatilis and the lens of roach Rutilus rutilus were identified on the basis of adults obtained by feeding various piscivorous birds of the families Laridae, Anatidae and Phasianidae with metacercariae on an experimental basis. Both morphological studies of the metacercariae and adults obtained from the intestine of the infected birds, as well as the suitability in these hosts, indicated that we were dealing with two different species of Diplostomum. The perch form was recognised as D. baeri Dubois, 1937. Although metacercariae of this species matured both in herring gull Larus argentatus and common tern Sterna hirundo, the principal final host seemed to be goosanders Mergus merganser. In contrast, the lens form in roach was identified as D. spathaceum (Rudolphi, 1819) and it seemed to be restricted to birds of the family Laridae. These experimental results agree with parasitological findings from birds infected in the wild.     

Birds and their food resources at Christchurch International Airport,New Zealand

The diversity and seasonal abundance of birds at Christchurch Airport, their food and feeding habits, and the phenology of the food resource were studied in 1968–69 as part of an investigation of the bird hazard to aircraft. The birds either fed and roosted on the airport or flew over or close to it. Seasonal trends in the diet of black‐backed gulls (Larus dominicanus), black‐billed gulls (L. bulleri), starlings (Sturnus vulgaris), and magpies (Gymnorhina tibicen hypoleuca) were determined by examining the gizzard contents of birds shot while they were feeding on the airfield. Marked food preferences were noticeably lacking, except that black‐backed gulls fed exclusively on earthworms. Black‐billed gulls, starlings, and magpies took a variety of insects, earthworms, arachnids, and, to a less extent, seeds. Species diversity, seasonal distribution, and abundance of invertebrates and seeds showed that the birds were opportunist feeders on a succession of temporarily abundant food, and their presence on the airfield was determined largely by the availability of invertebrates and seeds.     

Diel activity pattern and predation rate of the generalist predator Dicyphus hesperus

Abstract This study examined the diel activity pattern and the effect of diel activity pattern on predation rate and prey finding of Dicyphus hesperus Knight (Heteroptera: Miridae). To determine the diel activity pattern of D. hesperus, starved females were placed on tomato leaflets Lycopersicon esculentum Mill. (Solanaceae) under zero, low, or high light intensities at 02:00, 08:00, and 14:00 h, respectively, and the amount of time spent walking or resting during a 30‐min interval was recorded. Predation rates of D. hesperus females on Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) egg patches were determined under either a L16:D8 (long day) or L8:D16 (short day) diel period. Egg patches were removed from D. hesperus females after either 8 or 16 h of dark or 16 or 8 h of light, and the number of eggs consumed was counted. Dicyphus hesperus females spent more time searching for prey at night than during the day. Females ate eggs at a higher rate during the night than during the day. Overall, D. hesperus females had higher predation rates when reared under a long day diel cycle compared with females reared under a short day diel cycle. More females reared at the L16:D8 diel cycle found the egg patch during the night than during the day. There was no difference in egg patch finding between night and day for females reared at L8:D16. Overall, L16:D8 reared females found more egg patches than females reared at L8:D16. Therefore, D. hesperus females are more active and find and consume prey at a higher rate at night than day.     

Trophic niche width and overlap of two sympatric gulls in the southwestern mediterranean

The diets of two potential competitor species, Audouin's Larus audouinii and yellow-legged gulls Larus cachinnans, were examined while they bred at the Chafarinas Islands during 1993, 1994, and 1995. Data were collected during two commercial fishing regimes: (1) trawling and purse seine fisheries, and (2) diurnal trawlers only. Since the food supply for the gulls in this area was heavily reliant on the activity of purse seine fisheries, these contrasting situations allowed us to analyze short-term effects, induced by daily changes in food supply, on niche width, dietary shift, and niche overlap between the two species. Overall, both species relied mainly on fish for food, especially Clupeiforms, in the case of Audouin's gull irrespective of the fishing situation, and in the case of the yellow-legged gull, only when purse seine fishing was in operation. When purse seine boats did not operate (food shortage), yellow-legged gulls broadened their niche, consuming equal amounts of all the feeding resources, and they showed a dietary shift toward a greater consumption of prey from refuse tips. In contrast, Audouin's gulls did not change their niche width, but showed a slight dietary shift away from the consumption of epipelagic fish, compensated by an increase in reliance on benthic-mesopelagic resources. Niche overlap was clearly higher on days when both fishing fleets operated, probably because a superabundant food resource facilitates high overlap without affecting coexistence between the two species. Since our study was developed on the basis of daily variations in food supply, and competition effects are to be expected on a longer-term basis, these changes can be seen as the outcome of the coexistence of two species in stable competitive equilibrium. Received: 4 October 1996 / Accepted: 20 May 1997     

Food selection, food-seeking patterns and hunting success of captive Goosanders Mergus merganser and Red-breasted Mergansers M. serrator in relation to the behaviour of their prey

Food selection and food-seeking behaviour of hand-raised Goosanders Mergus merganser and Red-breasted Mergansers M. serrator were tested in experimental situations. When different combinations of two species of fish were presented simultaneously to hungry birds in a small trough where the prey was easily and equally available, neither duck species showed any preferences in prey selection. However, when these same fish species were presented to satiated birds, they showed definite preferences; the most preferred species were baltic salmon and brown trout followed in decreasing order by minnow, whitefish, sculpin, burbot and river lamprey. This was also the case when combinations of five to seven species were presented. However, under semi-natural conditions in a stream tank, the number of each species offish caught was related to their escape behaviour, not to the previously determined predator preferences. When salmon and minnows of different size were presented to hungry birds, they selected the larger fish regardless of species. In contrast, when satiated, they preferred small fish over large fish. Under the experimental conditions, the use of certain elements of foraging behaviour clearly differed between the two bird species: Mergus serrator spent more time hunting underwater than did M. merganser. Results are discussed in relation to the predator-prey situation existing under natural conditions in northern Swedish rivers.     

Diel variations in the assemblage structure and foraging ecology of larval and 0+ year juvenile fishes in a man‐made floodplain waterbody

This study investigated diel variations in zooplankton composition and abundance, and the species composition, density, size structure, feeding activity, diet composition and prey selection of larval and 0+ year juvenile fishes in the littoral of a man‐made floodplain waterbody over five 24 h periods within a 57 day period. There was a significant difference in the species composition of diurnal and nocturnal catches, with most species consistently peaking in abundance either during daylight or at night, reflecting their main activity period. There were no consistent diel patterns in assemblage structure or the abundance of some species, however, most likely, respectively, due to the phenology of fish hatching and ontogenetic shifts in diel behaviour or habitat use. There were few clear diel patterns in the diet composition or prey selection of larval and 0+ year juvenile roach Rutilus rutilus and perch Perca fluviatilis, with most taxa consistently selected or avoided irrespective of the time of day or night, and no obvious shift between planktonic and benthic food sources, but dietary overlap suggested that interspecific interactions were probably strongest at night. It is essential that sampling programmes account for the diel ecology of the target species, as diurnal surveys alone could produce inaccurate assessments of resource use. The relative lack of consistent diel patterns in this study suggests that multiple 24 h surveys are required in late spring and early summer to provide accurate assessments of 0+ year fish assemblage structure and foraging ecology.     

The predatory behavior of wintering <Emphasis Type="Italic">Accipiter</Emphasis> hawks: temporal patterns in activity of predators and prey

Studies focused on how prey trade-off predation and starvation risk are prevalent in behavioral ecology. However, our current understanding of these trade-offs is limited in one key respect: we know little about the behavior of predators. In this study, we provide some of the first detailed information on temporal patterns in the daily hunting behavior of bird-eating Accipiter hawks and relate that to their prey. During the winters of 1999–2004, twenty-one sharp-shinned hawks (A. striatus) and ten Cooper’s hawks (A. cooperii) were intensively radio tracked in rural and urban habitats in western Indiana, USA. Cooper’s hawks left roost before sunrise and usually returned to roost around sunset, while sharp-shinned hawks left roost at sunrise or later and returned to roost well before sunset. An overall measure of Cooper’s-hawk-induced risk (a composite variable of attack rate and activity patterns) generally reflected the timing of prey activity, with peaks occurring around sunrise and sunset. In contrast, risk induced by the smaller sharp-shinned hawk did not strongly reflect the activity of their prey. Specifically, an early morning peak in prey activity did not correspond to a period with intense hawk activity. The lack of early morning hunting by sharp-shinned hawks may reflect the high risk of owl-induced predation experienced by these hawks. The net effect of this intraguild predation may be to “free” small birds from much hawk-induced predation risk prior to sunrise. This realization presents an alternative to energetics as an explanation for the early morning peak in small bird activity during the winter.     

Habitat utilisation, feeding tactics and age related feeding efficiency in the Caspian Gull Larus cachinnans

The feeding behaviour of the Caspian Gull Larus cachinnans was analysed in southern Poland in 2001. During the pre-breeding period, most birds foraged on a refuse dump and some foraged in a river valley. During incubation, similar numbers of birds foraged on fishponds, gravel pits and the refuse dump. During the chick-rearing period, fishponds were the most important foraging grounds. The foraging success of three main foraging tactics was analysed: digging on refuse, fishing and kleptoparasitism. We found that digging success was higher in juveniles than in immature or adult birds. However, older birds moved and ate more items per unit of time than juveniles, which indicates that older birds improved their energy gain simply by a higher speed of searching. The opposite was found for fishing success. As juvenile birds made fewer attempts than immature or adult birds, fishing success was higher in adults. Adults and immature birds interrupted more attacks than juveniles, which indicates that older birds were better able to assess the probability of fish catching than juveniles. Kleptoparasitism was observed almost exclusively on the refuse dump during the pre-breeding period. Young birds kleptoparasitised more frequently than adults, but they had a lower rate of success. However, the lower success in young birds was due to victim choice, rather than differences in flight skills. Young birds kleptoparasitised Black-headed Gulls Larus ridibundus and Jackdaws Corvus monedula more frequently than adults, but none of the attacks towards these species was successful. Generally, Caspian Gulls kleptoparasitised conspecifics more often than expected from species frequency. Only attacks towards conspecifics yielded any success.     

Polymorphic microsatellite markers for the goosander (Mergus merganser)

To investigate the population genetic structure of the goosander (Mergus merganser) in Europe and to identify populations with a significant conservation value, we isolated nine microsatellite loci, and screened them in the subspecies Mergus merganser merganser and Mergus merganser americanus. All markers were polymorphic with two to 15 alleles per locus. Average observed and expected heterozygosity values were 0.422 and 0.624, respectively, for the European and 0.497 and 0.667, respectively, for the North American subspecies. Only one marker departed significantly from Hardy–Weinberg expectations in both subspecies. This marker was highly variable but homozygous in all females, suggesting a sex‐linked inheritance.     

Spatial contraction in a large gull colony in relation to the position of arctic fox dens

Many species of colonial ground-nesting birds are known to be sensitive to predation by terrestrial predators. The response of prey under the pressure of predation can either be direct (depletion of prey) or indirect due to prey avoiding the predator. We studied the recolonization of arctic foxes into a large and growing breeding colony of lesser black-backed gulls. The gull colony reached the size of 20,000 pairs during the period of no foxes. The number of breeding gulls continued to increase after the colonization of foxes and reached a maximum of 40,000 pairs about 15 years later. During the same time period, the spatial coverage of the gull colony shrunk from 31.4 down to 18.6 km², and the change in distribution was closely linked with position of fox dens, which ranged in number from one to three annually. In 2005, the number of breeding gulls decreased slightly with an ongoing shrinkage in spatial coverage. Food analyses from fox scats and stomachs showed that birds of the order Charadriiformes, particularly gulls, were the key prey item. Survival rates of gull nests were higher in the middle of the colony than at the colony edge close to a fox den. The colony area lost could be explained solely by the number of eggs removed by foxes but is unlikely, e.g., due to reclutching. We suggest that intraspecific predation contributes additionally to the effects of direct fox predation although to an unknown degree.     

At–Sea Behavior Varies with Lunar Phase in a Nocturnal Pelagic Seabird,the Swallow-Tailed Gull

Strong and predictable environmental variability can reward flexible behaviors among animals. We used long-term records of activity data that cover several lunar cycles to investigate whether behavior at-sea of swallow-tailed gulls Creagrus furcatus, a nocturnal pelagic seabird, varied with lunar phase in the Galápagos Islands. A Bayesian hierarchical model showed that nighttime at-sea activity of 37 breeding swallow-tailed gulls was clearly associated with changes in moon phase. Proportion of nighttime spent on water was highest during darker periods of the lunar cycle, coinciding with the cycle of the diel vertical migration (DVM) that brings prey to the sea surface at night. Our data show that at-sea behavior of a tropical seabird can vary with environmental changes, including lunar phase.     

Contrasting patterns of wetland use by sympatric larids in winter

The population dynamics and behaviour of the larid assemblage of a Mediterranean coastal wetland, the Vourkari inlet in Greece, were studied during the winter of 2008–2009. More black-headed gulls (Larus ridibundus) were seen in the inlet in December, more Mediterranean gulls (Larus melanocephalus) were present from mid-January to mid-February, while little variation was observed in yellow-legged gull (Larus michahellis) numbers throughout winter. Bird numbers remained stable through the day for the yellow-legged gull, but fewer black-headed and Mediterranean gulls were present in the late morning than the other periods. Diurnal activity patterns showed that feeding was the primary behaviour for yellow-legged gulls and resting and sleeping for black-headed and Mediterranean gulls, with locomotion being equally important for all species. All larids were primarily feeding in the late morning period, but yellow-legged gulls were doing so in much higher proportions. These contrasting patterns suggested that the Vourkari inlet was more important as a day roost for black-headed and Mediterranean gulls and a preferred feeding ground for yellow-legged gulls. Furthermore, results suggested that resting and sleeping were interchangeable activities and all other activities had more or less the same time demands on a daily basis, and also a consistent pattern across species in sleeping proportions that might indicate cross-species synchrony in sleep patterns; however future research is needed to resolve these issues. This study provided important new information on the winter ecology of three larids and revealed patterns of wetland use by these species that could help assess the importance of certain areas and improve coastal habitat management strategies to benefit birds.