Activity patterns of giant petrels, Macronectes spp., using different foraging strategies

We studied foraging activity of giant petrels during the incubation period, by simultaneously deploying activity recorders and satellite transmitters on northern (Macronectes halli) and southern giant petrels (Macronectes giganteus) at Bird Island (South Georgia, Antarctica) between 29 October and 26 December 1998. Satellite tracking showed two types of trips: (1) coastal trips, all undertaken by male northern giant petrels, to the nearby South Georgia mainland, presumably foraging on seal and penguin carcasses on beaches, and (2) pelagic trips, foraging at sea for marine prey or potentially scavenging on distant archipelagos (e.g. South Sandwich, Falkland or South Orkney Islands). Activity recorder data were consistent with the types of trip defined by the satellite tracking data, with median wet activity (time spent at the sea surface) during pelagic trips being 41%, but only 14% on coastal trips. On pelagic trips, there was a significant negative correlation between the duration of wet periods and the speed of travel between satellite uplinks. Mean travelling speed between uplinks was greater during day than night for both types of trips, suggesting that giant petrels prefer to travel during daylight and are less active at night. The scarcity of wet periods during the night in giant petrels foraging to the South Georgia coast (median=3%, range=1-9%) indicates that such birds spent almost all night on land. Likewise, the scarcity of wet periods at night for three birds foraging 700-1,000 km south of Bird Island, where there is no land but abundant icebergs, suggests these birds were resting on the icebergs at night. In addition to the adaptations to scavenging on carrion, pelagic trips by giant petrels contain elements similar to those of albatrosses, indicating a complexity to giant petrel lifestyle hitherto unrecognised.     

Do activity costs determine foraging tactics for an arctic seabird?

How energy costs affect foraging decisions is poorly understood for marine animals. To provide data relevant to this topic, we examined the relationship between activity levels and foraging behavior by attaching activity recorders to 29 chick-rearing wing-propelled diving birds (thick-billed murres, Uria lomvia) in 1999–2000. We connected the activity during the final dive bout with the prey item we observed being fed to the chicks. After accounting for changes in activity level with depth, activity was highest during the final dive of a dive bout, reflecting maneuvring during prey capture. Pelagic prey items, especially invertebrates (amphipods), were associated with higher depth-corrected activity, leading to shorter dives for a given depth (presumably due to higher oxygen consumption rates) and, thus, shorter search times (lower bottom time for a given depth). Pelagic prey items were likely captured during active pursuit, with the birds actively seeking and pursuing schooling mid-water prey. In contrast, benthic prey involved low activity and extended search times, suggesting that the birds slowly glided along the bottom in search for prey hidden in the sediments or rocks. We concluded that activity levels are important in determining the foraging tactics of marine predators. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Inter-annual changes in diet and foraging trip lengths in a small pelagic seabird, the thin-billed prion Pachyptila belcheri

Central place foragers are constrained in their foraging distribution by the necessity to return to their nest site at regular intervals. In many petrels that feed on patchily distributed prey from the sea surface over large foraging areas, alternating long and short foraging trips are used to balance the demands of the chick with the requirements of maintaining adult body condition. When the local conditions are favourable for prey density and quality, adults should be able to reduce the number of long foraging trips. We studied the flexibility in foraging trip lengths of a small pelagic petrel, the thin-billed prion Pachyptila belcheri, over three breeding seasons with increasingly favourable, cold-water conditions. During a warm-water influx in February 2006, chicks were fed less frequently and adults carried out foraging trips of up to 8 days. When conditions became more favourable with colder water temperatures in 2007 and 2008, thin-billed prions decreased trip lengths, more often attended their chick every day, and long foraging trips of six to eight days were not registered during 2008. Chick growth rates mirrored this, as chicks grew poorly during 2006, intermediate during 2007 and best during 2008. Thin-billed prions preyed mainly on squid during incubation and mainly on amphipods and euphausiids during chick-rearing. In the poorest season only, the diet was substantially supplemented with very small copepods. Together, the present results indicate that during warm-water conditions, thin-billed prions had difficulties in finding sufficient squid, amphipods or euphausiids and were forced to switch to lower trophic level prey, which they had to search for over large ocean areas.     

Trophic relationships of white-chinned petrels from Crozet Islands: combined stomach oil and conventional dietary analyses

The diet of white-chinned petrels Procellaria aequinoctialis breeding at the Crozet Archipelago (southern Indian Ocean) was studied using two complementary methods: lipid analysis of stomach oils as trophic markers together with the conventional dietary approach (i.e., stomach content analysis). Objectives were (1) to investigate the adult diet when they feed for themselves by analyzing stomach oil lipids, and (2) to compare the lipid signature of chick and adult oils. Stomach oils mainly consisted of triacylglycerols (TAG), diacylglycerol-ethers (DAGE) and wax esters (WE) (66, 14 and 11%, respectively). The dietary origin of TAG and WE was evaluated by linear discriminant analyses with fatty acid and fatty alcohol fractions. Analyses evidenced that stomach oils did not originate from Antarctic krill, but instead from myctophid fish, thus demonstrating the importance of mesopelagic fish in the nutrition of adult petrels. This result was consistent with the identification of digested remains of myctophids recovered from adult stomach contents after long foraging trips. Large amounts of a rare lipid class, DAGE (up to 76% of total lipids), were identified in two stomach oils, together with fresh remains of the squid Gonatus antarcticus (99% by mass), suggesting that DAGE could have the potential to be trophic markers of cephalopods. Moreover, six oils probably originated from Patagonian toothfish, thus confirming strong interactions between white-chinned petrels and fisheries. Comparison between chick and adult stomach oils indicated no major differences in their biochemical composition suggesting an identical dietary origin of oils, mainly myctophids. Both adult and chick oils can therefore be used to determine the feeding ecology of adult birds when they feed far away from their breeding grounds. Finally, food analysis of chick samples and adult samples collected after short and long trips indicated different foraging grounds during the two kinds of trips, and also between long trips performed in subtropical and Antarctic waters.     

Central place foraging by breeding Cook’s petrel Pterodroma cookii: foraging duration reflects range, diet and chick meal mass

Pelagic seabirds are central place foragers during breeding and variation in foraging trip duration and range reflect differences in diet and chick provisioning, through the exploitation of divergent habitats of varying productivity. We tested whether these relationships hold in small procellarriids by equipping chick-rearing Cook’s petrel Pterodroma cookii (200 g) with geolocation-immersion loggers, conducting isotope analysis of blood and measuring chick meal mass following foraging trips of varying duration. Cook’s petrel tracked during chick rearing from Little Barrier Island (LBI) and Codfish Island (CDF), New Zealand had larger maximum ranges during longer foraging trips. Blood nitrogen isotope signatures (δ¹⁵N) of adults were significantly higher after foraging trips of longer duration, but not of greater maximum range. There was no significant relationship between blood carbon isotope signatures (δ¹³C) and foraging trip characteristics. Proportion of time spent on the sea surface and the mass of the meal brought back to chicks were consistently greater for Cook’s petrel with larger maximum ranges, which in the case of birds from CDF coincided with productive subtropical convergence zone habitats. As predicted, trip duration reflected divergent foraging behaviours in Cook’s petrel during breeding. We suggest that the availability of different prey is a key factor governing at-sea distributions and dietary composition of this species.     

Contrasting foraging tactics by northern gannets (<Emphasis Type="Italic">Sula bassana</Emphasis>) breeding in different oceanographic domains with different prey fields

In order to forage and to provision offspring effectively, seabirds negotiate a complex of behavioural, energetic, environmental and social constraints. In first tests of GPS loggers with seabirds in North America, we investigated the foraging tactics of free-ranging northern gannets (Sula bassana) at a large and a medium-sized colony that differed in oceanography, coastal position and prey fields. Gannets at Low Arctic colony (Funk Island) 50 km off the northeast coast of Newfoundland, Canada provisioned chicks almost entirely with small forage fish (capelin Mallotus villosus, 89%), while at boreal colony (Bonaventure Island) 3 km from shore in the Gulf of St. Lawrence, Quebec, Canada, large pelagic fish dominated parental prey loads (Atlantic mackerel Scomber scombrus 50%, Atlantic herring Clupea harengus 33%). Mean foraging range and the total distance travelled per foraging trip were significantly greater at the larger inshore colony (Bonaventure) than at the smaller offshore colony (Funk Island; 138 and 452 km vs. 64 and 196 km, respectively). Gannets from Funk Island consistently travelled inshore to forage on reproductive capelin shoals near the coast, whereas foraging flights of birds from Bonaventure were much more variable in direction and destination. Birds from the Low Arctic colony foraged in colder sea surface water than did birds from the boreal colony, and dive characteristics differed between colonies, which is concordent with the difference in prey base. Differences between the colonies reflect oceanographic and colony-size influences on prey fields that shape individual foraging tactics and in turn generate higher level colony-specific foraging “strategies”.     

Persistent bimodal activity patterns in wild and captive black-tailed godwit <Emphasis Type="Italic">Limosa limosa</Emphasis> under different environmental conditions

There is a large literature dealing with daily foraging routines of wild birds during the non-breeding season. While different laboratory studies have showed that some bird activity patterns are a persistent property of the circadian system, most of field studies preclude the potential role of an endogenous circadian rhythm in controlling bird’s foraging routines. In this study we compared the patterns of diurnal foraging activity and intake rates of migrating black-tailed godwits, Limosa limosa (radio-tagged and non-tagged individuals) at two stopover sites (habitats) with different environmental characteristics, aiming at identifying proximate factors of bird activity routines. To gain insights into the role of food availability in control of such foraging routines, we also estimated foraging activity patterns in captive godwits subjected to constant food availability. Captive and wild black-tailed godwits showed a persistent bimodal activity pattern through daylight period. Food availability had a significant effect on the intake rates, but had a subtler effect on foraging and intake rate rhythms. Temperature and wind speed (combined in a weather index) showed non-significant effects on both rhythms. Although we could not discard a role for natural diurnal changes in light intensity, an important timing cue, our findings support the idea that an endogenous circadian rhythm could be an important proximate factor regulating foraging activity and food items taken per unit time of wild black-tailed godwits during migration.     

Pattern-oriented modeling of bird foraging and pest control in coffee farms

We develop a model of how land use and habitat diversity affect migratory bird populations and their ability to suppress an insect pest on Jamaican coffee farms. Bird foraging—choosing which habitat patch and prey to use as prey abundance changes over space and time—is the key process driving this system. Following the “pattern-oriented” modeling strategy, we identified nine observed patterns that characterize the real system's dynamics. The model was designed so that these patterns could potentially emerge from it. The resulting model is individual-based, has fine spatial and temporal resolutions, represents very simply the supply of the pest insect and other arthropod food in six habitat types, and includes foraging habitat selection as the only adaptive behavior of birds. Although there is an extensive heritage of bird foraging theory in ecology, most of it addresses only the individual level and is too simple for our context. We used pattern-oriented modeling to develop and test foraging theory for this across-scale problem: rules for individual bird foraging that cause the model to reproduce a variety of patterns observed at the system level. Four alternative foraging theories were contrasted by how well they caused the model to reproduce the nine characteristic patterns. Four of these patterns were clearly reproduced with the “null” theory that birds select habitat randomly. A version of classical theory in which birds stay in a patch until food is depleted to some threshold caused the model to reproduce five patterns; this theory caused lower, not higher, use of habitat experiencing an outbreak of prey insects. Assuming that birds select the nearby patch providing highest intake rate caused the model to reproduce all but one pattern, whereas assuming birds select the highest-intake patch over a large radius produced an unrealistic distribution of movement distances. The pattern reproduced under none of the theories, a negative relation between bird density and distance to trees, appears to result from a process not in the model: birds return to trees at night to roost. We conclude that a foraging model for small insectivorous birds in diverse habitat should assume birds can sense higher food supply but over short, not long, distances.     

Sea-bird aggregation at a deep North Pacific seamount

During June 1991, we studied sea birds at a mid-ocean seamount (Fieberling Guyot) in the eastern North Pacific Ocean. Avifaunal composition changed from small Procellariiformes [a storm-petrel; Oceanodroma leucorhoa (Vieillot)] away from the seamount to an assemblage dominated by larger tubenoses [mostly black-footed albatross Diomedea nigripes Audubon and Cook's petrel Pterodroma cookii (Gray)]. Compared to adjacent waters, sea-bird density and biomass within a 30-km radius centered on the seamount summit were 2.4 and 8 times higher, respectively. Individual sea-bird taxa were 2 to 40 times more abundant at the seamount relative to values reported previously from large-scale surveys of deep-ocean regions in the central North Pacific. In September 1991 we studied potential prey of sea birds in the upper water column using a neuston net and multiple opening-closing net system (MOCNESS) tows. Most potential prey types in the neuston exhibited no significant enhancement over the seamount. MOCNESS samples at 10 m depth, however, showed several prey types to be more abundant over the seamount, and the dominant size class of fish was slightly larger. We attribute the sea-bird aggregation observed at this seamount to changes in the abundance and/or behavior of pelagic organisms in the deep scattering layer (not adequately sampled in this study), perhaps augmented by migrations of seamount residents into the surface layers. Processes on and in the vicinity of seamounts may provide spatially-predictable prey to wide-ranging aerial sea birds foraging in this relatively austere environment.     

Primary versus secondary drivers of foraging activity in sandeel schools (<Emphasis Type="Italic">Ammodytes tobianus</Emphasis>)

The commercially and ecologically valuable sandeel (Ammodytes ssp.) make distinct vertical shifts between an inactive stage, during which they seek refuge in the sand, and a pelagic schooling stage, during which they forage. This characteristic discontinuous foraging pattern constitutes a challenge to fishery biologists and has consequences for a wide range of predators ranging from birds and mammals to commercially important species. However, experimental studies that shed light on the primary drivers of foraging activity in fish are rare. In the present study, whole schools of sandeel (A. tobianus) were caught in August in east Denmark (65°02′30N; 12°37′00E) and kept in large tanks in the laboratory. It was found that the amount of food ingested and memory of past days feeding history are primary drivers of foraging activity at the level of the entire school, whereas external factors such as prey concentration and temperature are merely secondary drivers.     

Diurnal and nocturnal feeding rate in Kentish plovers <Emphasis Type="Italic">Charadrius alexandrinus</Emphasis> on an intertidal flat as recorded by telescopic video systems

To elucidate the effects of light, site, temperature, time after emersion, and wind speed on foraging attempt rate (FAR) (attempts/unit time) and feeding success (FS) (captures/attempts) in the obligate visual foraging shorebird, Kentish plovers Charadrius alexandrinus, field observations were performed at a sandflat in Tokyo Bay, using a visible-light telescope camcorder during the day and a thermal infrared telescope camcorder at night. The re-analysis capability and frame-step replay of highly magnified video-images can contribute to accurate measurement of feeding behaviour over conventional focal observation techniques. Kentish plovers increased both FAR and FS in areas of high prey (polychaetes and crabs) density, resulting in a synergistic increase in feeding rate (captures/unit time). In areas of high prey abundance, FAR was higher at lower wind speed. FS increased with increasing time after emersion. Increasing temperatures induced a positive effect on FAR but a negative effect on FS. The effect of light on FAR was not observed; however, time-to-defecation occurrence was 2.2-fold shorter at night than during the day, indicating that feeding rate and FS are higher at night. These results are attributable to an increase in availability of cues due to higher nocturnal activity in polychaetes. Since available foraging time (emersion time) at night was 1.7-fold longer than during the day in the present study period, Kentish plovers could capture 3.7-fold (2.2 × 1.7) more prey at night than during the day. Results from this study indicate that nocturnal feeding in overwintering plovers is not a compensation but a major foraging activity to meet their energy requirements. An erratum to this article can be found at     

What limits predator detection in blue tits (<Emphasis Type="Italic">Parus caeruleus</Emphasis>): posture,task or orientation?

To detect threats and reduce predation risk prey animals need to be alert. Early predator detection and rapid anti-predatory action increase the likelihood of survival. We investigated how foraging affects predator detection and time to take-off in blue tits (Parus caeruleus) by subjecting them to a simulated raptor attack. To investigate the impact of body posture we compared birds feeding head-down with birds feeding head-up, but could not find any effect of posture on either time to detection or time to take-off. To investigate the impact of orientation we compared birds having their side towards the attacking predator with birds having their back towards it. Predator detection, but not time to take-off, was delayed when the back was oriented towards the predator. We also investigated the impact of foraging task by comparing birds that were either not foraging, foraging on chopped mealworms, or foraging on whole ones. Foraging on chopped mealworms did not delay detection compared to nonforaging showing that foraging does not always restrict vigilance. However, detection was delayed more than 150% when the birds were foraging on whole, live mealworms, which apparently demanded much attention and handling skill. Time to take-off was affected by foraging task in the same way as detection was. We show that when studying foraging and vigilance one must include the difficulty of the foraging task and prey orientation.Communicated by P.A. Bednekoff     

Latitudinal clines in food distribution correlate with differential migration in the Western Sandpiper

We report that a latitudinal cline in intertidal food distribution is associated with the nonbreeding distribution of the Western Sandpiper (Calidris mauri). This novel result is the first to demonstrate a clear relationship between patterns of differential nonbreeding distribution and food availability for any shorebird species. Within each age class and sex, longer-billed Western Sandpipers winter further south. Moreover, females, the longer-billed sex, tend to winter south of males. Thus, both inter- and intra-sexual clines in bill morphology result in an overall gradient of increasing bill length from north to south. Longer-billed birds are able to extract prey that are buried more deeply in the sediment; therefore, we predicted shifts in the vertical distribution of food resources to coincide with the clines in bill morphology across the nonbreeding range. We tested our prediction by measuring biofilm density and the vertical distribution of macrofaunal invertebrates at six nonbreeding sites. Although no latitudinal trend was observed for biofilm, the vertical distribution of invertebrates was consistent with our prediction and revealed that the greatest relative abundance of surface prey occurred at northern nonbreeding sites and declined with decreasing latitude. We discuss the potential implications of these findings in the context of competing evolutionary hypotheses of differential migration and bill dimorphism in shorebirds.     

<Emphasis Type="Italic">Octopus insularis</Emphasis> (Octopodidae), evidences of a specialized predator and a time-minimizing hunter

Shallow-water octopuses have been reported as major predators of motile species in benthonic marine communities, capturing their prey by different foraging techniques. This study assessed for the first time the feeding ecology, foraging behavior, and defensive strategy during foraging, including the use of body patterns, to construct a general octopus foraging strategy in a shallow water-reef system. Octopus insularis was studied in situ using visual observations and video recordings. The diet included at least 55 species of crustaceans (70%), bivalves (17.5%), and gastropods (12.5%); however, only four species accounted for half of the occurrences: the small crabs Pitho sp. (26.8%) and Mithrax forceps (23.9%), the bivalve Lima lima (5.3%), and the gastropod Pisania pusio (4.9%). Poke and crawl were most frequent foraging behaviors observed in the video recordings. The foraging behaviors were associated with environmental variables and octopus body size. The sequences of foraging behavior showed characteristics of a tactile saltatory searching predator, as well as a visual opportunist. Body patterns showed a relationship with foraging behavior, habitat variables, and octopus body size. Mottle was the most frequent pattern, especially during poke and crawl, in shallower depths. Dorsal light–ventral blue green was more frequent during swimming at mid-water, and Blotch was the normal pattern during web-over by large animals. The large proportion of two species of small crabs in den remains, the intense search for food during short hunting trips, and the intense use of cryptic body patterns during foraging trips, suggest that this species is a ‘time-minimizing’ forager instead of a ‘rate-maximizer’.     

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

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

Time stress,predation risk and diurnal–nocturnal foraging trade-offs in larval prey

Insect larvae increase in size with several orders of magnitude throughout development making them more conspicuous to visually hunting predators. This change in predation pressure is likely to impose selection on larval anti-predator behaviour and since the risk of detection is likely to decrease in darkness, the night may offer safer foraging opportunities to large individuals. However, forsaking day foraging reduces development rate and could be extra costly if prey are subjected to seasonal time stress. Here we test if size-dependent risk and time constraints on feeding affect the foraging–predation risk trade-off expressed by the use of the diurnal–nocturnal period. We exposed larvae of one seasonal and one non-seasonal butterfly to different levels of seasonal time stress and time for diurnal–nocturnal feeding by rearing them in two photoperiods. In both species, diurnal foraging ceased at large sizes while nocturnal foraging remained constant or increased, thus larvae showed ontogenetic shifts in behaviour. Short night lengths forced small individuals to take higher risks and forage more during daytime, postponing the shift to strict night foraging to later on in development. In the non-seasonal species, seasonal time stress had a small effect on development and the diurnal–nocturnal foraging mode. In contrast, in the seasonal species, time for pupation and the timing of the foraging shift were strongly affected. We argue that a large part of the observed variation in larval diurnal–nocturnal activity and resulting growth rates is explained by changes in the cost/benefit ratio of foraging mediated by size-dependent predation and time stress.     

Prey-capture benefits in a mixed-species group of Amazonian tamarins,Saguinus fuscicollis and S. mystax

Summary Tamarins of the genus Saguinus feed on a wide range of arthropods and small vertebrates, which compose a critical component of their diet. This paper examines the foraging patterns and capture success of the Avila-Peres saddle-back (S. fuscicollis avilapiresi) and the red-capped moustached tamarin (S. mystax pileatus) in very stable mixed-species groups, and whether and how any foraging benefits for either species resulted from their association. Moustached tamarins actively searched for prey items which were mainly well exposed on the midstorey foliage. Saddle-back tamarins, on the other hand, foraged at lower heights, largely by manipulating a variety of microhabitats potentially concealing embedded prey. The foraging activity of the numerically dominant and larger-bodied moustached tamarins often resulted in prey items escaping to lower substrates, usually the forest leaf-litter. The beating effect of this species substantially facilitated captures of large, mobile prey items by saddle-backs, which were highly adept at locating and retrieving flushed prey. It is estimated that, while saddle-backs obtained 66–73% of their prey biomass from flushed items, this proportion was substantially lower (2–9%) for moustached tamarins. Commensal insectivory appears to involve a highly asymmetric benefit to saddle-backs, and a low cost to moustached tamarins, which partly explains the stability of mixed-species groups. Correspondence to the present address     

Shell ecophenotype in the blue mussel (<Emphasis Type="Italic">Mytilus edulis</Emphasis>) determines the spatial pattern in foraging behaviour of an oystercatcher (<Emphasis Type="Italic">Haematopus ostralegus</Emphasis>) population

When feeding on blue mussels (Mytilus edulis), oystercatchers (Haematopus ostralegus) either stab into the mollusc’s gaping valves or hammer through its dorsal or ventral shell. Whilst the selectivity of hammering and stabbing oystercatchers for specific prey morphologies has been well studied, the way in which the effects of environment on M. edulis morphology can in turn affect feeding methods of H. ostralegus is very poorly understood. Based on morphological analyses on randomly selected shells from three intertidal zones, this study failed to detect differences in morphology or distribution of dorsally and ventrally hammered shells but confirms the finding of previous authors that hammering oystercatchers select thinner mussels than stabbing birds. Additionally, we show that this difference in optimal prey morphology can lead to spatial patterns in oystercatcher feeding behaviour. Whilst at the low intertidal and higher mid intertidal zones, characterised by comparatively thick shells, most empty shells had apparently been stabbed, hammering was the dominant feeding behaviour at the lower mid intertidal zone, where shells were thinner. Preference of hammering birds for smaller mussels was not ubiquitous. Sagittal shell shape was predominantly influenced by allometric growth effects and had only minor effect on prey selection. All oystercatchers preferred less inflated mussels, with the degree of shell inflation gradually increasing with higher intertidal elevation. Our results illustrate the importance of small-scale patterns in prey ecophenotypes in determining the distribution and feeding dynamics of wading birds.     

Diurnal and nocturnal feeding rate in Kentish plovers <Emphasis Type="Italic">Charadrius alexandrinus</Emphasis> on an intertidal flat as recorded by telescopic video systems

To elucidate the effects of light, site, temperature, time after emersion, and wind speed on foraging attempt rate (FAR) (attempts/unit time) and feeding success (FS) (captures/attempts) in the obligate visual foraging shorebird, Kentish plovers Charadrius alexandrinus, field observations were performed at a sandflat in Tokyo Bay, using a visible-light telescope camcorder during the day and a thermal infrared telescope camcorder at night. The re-analysis capability and frame-step replay of highly magnified video-images can contribute to accurate measurement of feeding behaviour over conventional focal observation techniques. Kentish plovers increased both FAR and FS in areas of high prey (polychaetes and crabs) density, resulting in a synergistic increase in feeding rate (captures/unit time). In areas of high prey abundance, FAR was higher at lower wind speed. FS increased with increasing time after emersion. Increasing temperatures induced a positive effect on FAR but a negative effect on FS. The effect of light on FAR was not observed; however, time-to-defecation occurrence was 2.2-fold shorter at night than during the day, indicating that feeding rate and FS are higher at night. These results are attributable to an increase in availability of cues due to higher nocturnal activity in polychaetes. Since available foraging time (emersion time) at night was 1.7-fold longer than during the day in the present study period, Kentish plovers could capture 3.7-fold (2.2 × 1.7) more prey at night than during the day. Results from this study indicate that nocturnal feeding in overwintering plovers is not a compensation but a major foraging activity to meet their energy requirements.     

Parent-offspring aggression in moorhens

Summary Colonial orb-weaving spiders from Mexico were studied to test predictions of risksensitive foraging theory: 1. group foraging increases prey capture/individual, and reduces prey variance; 2. spiders should be expected to exhibit risk-averse behavior (forage in groups) when the average level of prey exceeds individual needs, and exhibit risk-prone behavior (forage solitarily) when prey are searce. Laboratory and field studies show that group foraging increases capture efficiency and reduces variability in prey captured per spider. In desert/mesquite grassland habitat, where prey availability is low, M. atascadero forage solitarily in most cases. In tropical rainforest/agriculture sites, M. increassata forage in large colonies of thousands of webs. In intermediate habitats, M. spinipes forages solitarily or in groups, depending on prey availability. Over a range of sites with varying levels of prey, M. spinipes shifts from a risk-prone to a risk-averse group foraging strategy as prey increases.Group foraging behavior observed in colonial Metepeira fits the predictions of risk-sensitive foraging models. These findings explain why spiders tend to group webs together only in areas of superabundant prey. The role of risk-sensitivity in the evolution of coloniality in spiders is discussed.