Individual differences in aggressiveness and food stealing among wintering oystercatchers, Haematopus ostralegus L

Oystercatchers feeding on mussels foraged in small, overlapping areas and aggressive encounters were common amongst all birds irrespective of their method of opening mussels. Individuals varied consistently in aggressiveness. The more aggressive birds were more frequently present on the mussel bed, fed in those parts where food was most abundant, and were more successful in stealing mussels from other birds. Only the more aggressive birds stole more mussels than they had stolen from them, and stealing increased their rate of food intake.     

Piping as a display of dominance by wintering Oystercatchers Haematopus ostralegus

Although Oystercatchers Haematopus ostralegus wintering in the Exe Estuary performed the piping display in any type of aggressive encounter, it was predominantly observed in two contexts. ‘Solitary piping’ occurred when one individual piped without an obvious opponent being in the immediate vicinity. In ‘piping ceremonies’, two or more individuals piped at each other for prolonged periods at close distance. Prolonged contests over mussels were usually won by the individual that had initially started piping. The most aggressive and successful birds (the dominants), engaged most often in solitary piping and piping ceremonies, using up to 10% of their time this way. This suggests that the behaviour was associated with dominance rank. The hypothesis is that the two birds are likely to start piping if their relative status has not already been established, so that a quick end to a dispute over a mussel or feeding site is not achieved. A dominant individual performs solitary piping from a distance to suppress the aggressive activities of other birds within its feeding range. If this signal fails, the dominant may join the ceremony, explaining why three was the most frequent number of birds involved in piping ceremonies. The ‘butterfly’ flight, used mainly by apparently dominant birds as they arrived on the feeding grounds, may signal their high status to birds already there. The outcome of several piping ceremonies involving birds of known dominance was consistent with this explanation. The location of piping ceremonies involving only two birds (interpreted as piping ceremonies not joined by dominants) at the extremities of the feeding ranges of the most dominant individuals is also consistent with this hypothesis, because this is where their dominance would be expected to decline and would therefore be more likely to be contested. This explanation of piping and butterfly displays assumes that individuals can recognize other individuals or quickly assess their dominance. Since dominant individuals were attacked less often than subdominants and attacking birds were rebuffed less than would be expected if recognition did not occur, we suggest that some form of dominance or individual recognition did exist. The width of the white collar did not correlate with dominance, but the individual variations in this feature may enable birds to distinguish each other and thus learn their relative status. It is intriguing that the diplomatist posture, which is typically used to rebuff misguided attackers, seems almost designed to give the opponent a particularly clear view of the white collar.     

Spatial consequences of aggressive behaviour in flocks of oystercatchers, Haematopus ostralegus L.

Oystercatchers were distributed non-randomly on mussel-beds in winter; the loose foraging flocks showed a ‘spaced-out gregariousness”. Aggressive defence of personal space within the flock was evidently not responsible for this spacing, as most attacks were not directed against birds unusually close to aggressors. Mutual avoidance, rather than overt aggression, apparently maintained the observed non-random spacing. Oystercatchers seemed to attack others in order to steal newly-opened mussels, but the likelihood of successfully gaining an illicit mussel in this way declined as the initial distance between attacker and victim increased. In this situation, the risk of falling victim to food theft may make avoidance of conspecifics advantageous.     

Individual and age differences in the feeding ecology of Oystercatchers Haematopus ostralegus wintering on the Exe Estuary, Devon

The habitats and foods used by a sample of marked individuals and by the whole population of Oystercatchers Haematopus ostralegus wintering on the Exe estuary, South Devon, are described. In the spring and early summer, only a few hundred immatures were present. Two thousand adults and several hundred juveniles arrived in late summer from the breeding grounds and remained until the following spring. The majority of the birds were then adults. At low water, most adults fed on the mussel Mytilus edulis beds and ate mussels. Most individuals specialised on this prey and ate little else. Some adults fed on mud-flats and sandflats within the estuary and along the coast, and specialised on a mixture of Nereis diversicolor and Scrobicularia plana or on Cerastoderma edule or Littorina spp. Though some juveniles ate mussels from the time they arrived, most did not. In winter, they took Scrobicularia on mudflats, earthworms Lumbricidae and leatherjackets Tipulidae larvae in fields and Spisula and mussels along the coast. At other times of year they mainly ate Nereis. However, as they grew older, more birds began to specialise on mussels, especially in their second, third and fourth summers when the adults were away. At high water, most birds roosted at the mouth of the estuary or along the coast. However, several hundred fed in the fields in winter with more doing so late in the winter on warm days on Neap tides when less time was available for feeding on the estuary in daylight. Birds feeding on Nereis and Scrobicularia at low water were most often seen in the fields, irrespective of age. Birds feeding on Mytilus and Littorina occurred there rarely, but no bird eating Cerastoderma was seen there. The findings are discussed in relation to age differences in feeding skills and ability to compete successfully with other birds.     

Locally stable wintering numbers in the Oystercatcher Haematopus ostralegus where carrying capacity has not been reached

Since 1976, the numbers of Oystercatchers Haematopus ostralegus wintering on the Exeestuary have fluctuated independently of the substantial increase over the same period in the numbers wintering in Britain as a whole. While this might imply that the carrying capacity of the Exe has been reached, closer examination suggests not. Over a sample of the years studied, Oystercatcher numbers on the main Mussel Mytilus edulis feeding areas increased even though the mussel bed quality remained unchanged, food abundance decreased and disturbance on some important beds increased. There was some redistribution of birds between mussel beds, apparently linked to the increased bird numbers and to a relative improvement in the food supply on two preferred mussel beds. The increase in Oystercatcher numbers on the main feeding areas over a period in which the feeding conditions did not improve suggests that carrying capacity was not reached on the mussel beds themselves and, therefore, on the estuary as a whole.     

The influence of predation by herring gullsLarus argentatus and oystercatchersHaematopus ostralegus on a newly established musselMytilus edulis bed in autumn and winter

Predation by herring gullsLarus argentatus and oystercatchersHaematopus ostralegus was evaluated on a newly established musselMytilus edulis bed on tidal flats of the German Wadden Sea. The mussel bed covered an area of 2 ha and showed a decrease in biomass of 40% in the most densely covered parts from August to January. Synchronously, the extent of the mussel bed was reduced, resulting in a decrease of average biomass of 98% over the whole mussel bed. From the beginning of August 1994 to mid January 1995, the average size of mussels increased from 10.7 to 20.3 mm. The P/B-ratio was 0.68 in August and 0.18 between September and November. Herring gulls and oystercatchers were the most important mussel predators. On average, 266 herring gulls and 63 oystercatchers were present on the mussel bed during one low tide; 34% of the herring gulls and 78% of the oystercatchers were observed to be feeding. Herring gulls fed at a rate of 4.2 mussels per minute and oystercatchers at a rate of 1.3 mussels per minute. While herring gulls took the most common mussel sizes (mean: 20 mm), oystercatchers searched for the largest mussels available (mean: 25 mm). Herring gulls consumed 13 mussels/m² (0.3g AFDW) during one day and oystercatchers 1.7 mussels/m² (0.1 g AFDW). Predation by birds was compensated by 33% of the production. The proportion removed by bird predation amounted to 10% of abundance and to 16% of biomass (including production). Oystercatchers were responsible for 1% of the reduction in abundance and for 3% of biomass. Removal was highest in the most common size classes of mussels, mainly caused by herring gulls. However, the highest proportion of mussels was eaten in the largest size classes, mainly by oystercatchers. *** DIRECT SUPPORT *** A03B6035 00004     

Disentangling interference competition from exploitative competition in a crab–bivalve system using a novel experimental approach

In predator–prey relationships such as those between crabs and their bivalve prey, interference competition is a topic of intense investigation as it can have profound consequences on the dynamics of both predator and prey populations. However in laboratory experiments – also those on crab–bivalve systems – workers never adequately disentangled interference competition from exploitative competition, as prey depletion was never compensated. Hitherto, experimental studies on crab–bivalve systems lack direct behavioural observations and have provided only indirect and thus inconclusive evidence of interference competition. We studied interference competition in adult male shore crabs Carcinus maenas that foraged on blue mussels Mytilus edulis. We developed a novel type of experimental tank to replenish each consumed mussel, and thus to keep prey levels constant. We conducted two experiments in which we varied number of crabs (1, 2, 4) and number of mussels (first experiment: 4, 8, 16, 32; second experiment: 8, 32, 128) and directly observed the foraging behaviour of crabs (foraging area=0.25 m²). In the first experiment, feeding rates decreased with increasing crab density only at mussel density 16 because both search time and time spent in agonistic interactions increased. At other mussel densities, variation in crab density did not affect feeding rates, possibly because of low statistical power and the narrow range of mussel densities offered. In the second experiment feeding rates decreased with increasing crab density because crabs spent more time in agonistic interactions and handling their prey. Feeding rates increased with increasing mussel density. Overall, crabs spent on average 14–18% of their foraging time in agonistic behaviours, while on three out of 64 occasions feeding rates decreased because mussels were stolen (kleptoparasitism). Concluding, we have shown that interference competition occurs in absence of prey depletion, while conducting direct behavioural observations aid to identify the behavioural processes that underlie interference competition.     

Feeding ecology of the Pacific Golden Plover (Pluvialis fulva) in the Sg. Labu River on the west coast of the Malay Peninsula

The feeding ecology of the Pacific Golden Plover (Pluvialis fulva) in the Sg. Labu river near the Matang Mangrove Forest Reserve on the west coast of the Malay Peninsula was studied in March and April of 1994. The plovers had two foraging sites, a polychaete-feeding and a mussel-feeding site, which were located at a distance of about 1 km from each other. The tidal height of the polychaete-feeding site was higher than that of the mussel-feeding site, which was not exposed at the neap tide. Therefore, the polychaete-feeding site, which was exposed during every low tide, was better for food collection than the mussel-feeding site. The individuals that undertook more frequent changes in feeding sites had a lower intake rate of polychaetes in the polychaete-feeding site. There were two classes in the feeding order, and the lower class had to feed on mussels in addition to polychaetes to obtain sufficient food.For polychaete-eaters, the total number of steps during a feeding bout was significantly higher and the number of pecks was significantly lower than those of mussel-eaters. Usually, mussel-eaters spent more time preening and resting than polychaete-eaters, although the former had less time for feeding than the latter had because of the shorter exposure time of the mussel-feeding site. This means that mussel-eaters may have had higher feeding efficiency than polychaete-eaters, regardless of their preference for polychaetes as a diet.Polychaete larvae frequently settled on mussel beds and their populations became established after the collapse of mussel populations. Therefore, mussels were not only an important part of the diet of P. fulva, but they also constructed mussel beds that attract polychaetes, i.e. the plover's most important food.     

The influence of flock size and geometry on the scanning behaviour of spotted turtle doves,Streptopelia chinensis

Abstract The negative correlation between the time individuals spend scanning the environment for predators and group size is usually explained by the benefit of corporate vigilance. However, this negative correlation may be explained additionally in terms of the ‘dilution effect’ and ‘selfish herd geometry’. Our experimental investigation of the scanning behaviour of free-living spotted turtle doves foraging at different shaped feeders revealed that flock geometry influenced individual scanning rates. The time spent scanning declined with group size less rapidly among birds foraging in linear flocks than among those foraging in more two-dimensional flocks. These results were not confounded by aggressive behaviour, and indicate that the benefits of foraging in groups include the so-called selfish herd geometry.     

Turbidity-mediated interactions between invasive filter-feeding mussels and native bioturbating mayflies

1. Invasive dreissenid mussels are known to cause large ecosystem changes because of their high filter‐feeding capacity, while native bioturbators may interfere with the mussels filter feeding. In this experiment, we investigated indirect environmental interactions between invasive filter‐feeding dreissenid mussels (zebra and quagga mussels) and native recolonizing bioturbating hexagenid mayflies (Hexagenia) at two mussel densities and two Hexagenia densities in a 2‐month long laboratory experiment. 2. Mean turbidity increased with increasing density of Hexagenia and decreased with increasing density of mussels. Turbidity showed the fastest decline at the highest mussel density, and no decline or a lower rate of decline at the low mussel density, dependent on Hexagenia density. 3. Mussel growth decreased with increasing Hexagenia density at low but not at high mussel density. Moreover, growth of mussels decreased as a function of increased mean turbidity at low mussel density but not at high mussel density. Filtering activity at the highest mussel density increased after introduction of food at the lower two densities of Hexagenia, but was constantly high at the highest Hexagenia density. 4. There was no difference in emergence of Hexagenia among the treatments, but mortality of Hexagenia was higher in the presence of mussels than in their absence. 5. Our results indicate that interactions between dreissenids and hexagenids are mediated through the sediment, and depend on density of both dreissenids and hexagenids. As the natural densities of these animals vary considerably within lakes, their growth and survival because of indirect environmental interactions is expected to vary spatially.     

SEASONAL CHANGES IN BODY-WEIGHT OF OYSTERCATCHERS HAEMATOPUS OSTRALEGUS

The body-weights of Oystercatchers Haematopus ostralegus wintering in Morecambe Bay, north-west England, showed marked seasonal changes between late summer and late winter, with considerable differences apparent between adult and immature birds. An attempt is made to relate these changes to recorded seasonal variations in prey biomass and to the annual cycles of breeding, moult and migration of the Oystercatcher. The mean weight of females invariably exceeded the mean weight of males in samples collected on the same dates, regardless of age. Adults returned from northern breeding areas in very lean condition, with mean weights ranging from 526 g in males to 540 g in females. Mean weight then increased progressively, due mainly to fat deposition, to a peak in March (up to 662 g in males and 675 g in females) around the time of their main departures for breeding. Heaviest birds then exceeded 800 g. Birds migrating to Iceland in spring would need to be of above average weight in March to make the shortest crossing (850 km, 13 h), via Scotland, while Oystercatchers of 700 g and over could probably make a direct flight (1500 km, 25 h) from Morecambe Bay in favourable weather. Breeding weights of British Oystercatchers were similar to those of post-breeders returning to Morecambe Bay in late August. The mean weights of first-year Oystercatchers arriving in August were very low, 449 g in males and 478 g in females. Their weights, and those of second- and third-year immatures, then rose rapidly in autumn, with some fat deposition, and reached mean values ranging between 551 g (males) and 597 g (females) by November-December. Mean weight then fell by 10–17% from December to March returning close to or below the September levels, whereas adults gained a further 6% during these winter months. Summer and autumn weight gains, and the major moult of adults and older immatures, occurred when the biomass of their two staple mollusc preys, Mytilus edulis (mussel) and Cardium edule (cockle), was maximal. Winter loss in mean weight of immatures corresponded with declining prey biomass, suggesting either that they were less efficient than adults in coping with deteriorating winter food supplies, or that they had no need to accumulate further (premigratory) fat reserves. The autumnal increases in mean weight of immatures are interpreted as an adaptation for withstanding adverse feeding conditions in winter. The Oystercatcher appears to be the only wader species in Britain in which adults increase, rather than lose, weight during the winter. This may be a consequence of an early breeding season, but it may be regarded also as a measure of the success Oystercatchers have achieved by specializing on a difficult but plentiful prey source.     

Phosphorus recycling by zebra mussels in relation to density and food resource availability

Using flow-through microcosms, we examined phosphorus (P) recycling by zebra mussels under conditions of nearly constant food resource supply and varying zebra mussel population densities (600–5200 ind./m²). At all density levels, zebra mussels filtered substantial algae, measured as chlorophyll biomass. Because chlorophyll biomass inputs were low throughout the study, zebra mussel biomass-specific rates of chlorophyll filtration declined with increasing density, suggesting food resource limitation at the higher densities. We observed net total P export and high zebra mussel biomass-specific rates of P recycling over time in microcosms at high zebra mussel densities. In systems with a low zebra mussel density, net total P export did not occur over time. Our results suggest the occurrence of P remineralization by zebra mussels and net loss associated with emaciation during periods of temporary starvation. These findings have implications for P dynamics since zebra mussels can be subjected to periods of starvation over seasonal and annual time scales.     

Reciprocal Effects between Burying Behavior of a Larval Dragonfly (Odonata: Macromia illinoiensis) and Zebra Mussel Colonization

Invasive zebra mussels (Dreissena polymorpha) often colonize dragonfly larvae, especially spawling species whose survivorship to emergence as terrestrial predators is consequently reduced. Using individuals of the sprawler, Macromia illinoiensis, as their own controls, we compared the burying behavior of penultimate instar larvae before (i.e. baseline) and after their colonization by zebra mussels under ambient conditions. Individuals that took longer to bury themselves when mussel-free had a higher rate of colonization by mussels over a five-day period compared to those that buried faster. In contrast, the depth at which individuals buried when mussel-free was not predictive of subsequent colonization rate. Although mean bury time did not differ between baseline and when an individual carried one or more mussels, colonized larvae buried more shallowly than when mussel-free. Moreover, attached mussels increased the risk of subsequent colonization by zebra mussels. After naturally losing all of their attached mussels, bury time and depth of individuals did not differ from their baseline behavior, indicating that the changes in the behavior of colonized individuals were due to mussel loads and not their time in captivity. Under natural conditions, the positive feed-back between mussel attachment and increasing vulnerability to colonization helps explain how mussel loads, which are lost at molting, can accumulate quickly over the duration of the final larval stadium. Because zebra mussel attachment decreases the crypsis that that a M. illinoiensis gains from burying, the invasive mussel may also make dragonfly larvae more detectable to visual predators.     

Effect of anthropogenic feeding regimes on activity rhythms of laboratory mussels exposed to natural light

Anthropogenic disturbance may affect animal behaviour and should generally be minimised. We examined how anthropogenic disturbance (24 h food deprivation) affected circadian rhythms in laboratory mussels Mytilus edulis exposed to natural light in the absence of tides. Repeated measures data were collected on mussel gape angle, exhalant pumping and valve adduction using a Hall sensor system over eight consecutive 24 h periods when exposed to two feeding conditions after 24 h food deprivation. Mussels (fed once per day at either midday or midnight) exposed to natural light showed a clear day–night rhythm with increased nocturnal activity: significantly greater gape angle, increased exhalant pumping and had significantly higher valve adduction rates. However, circadian rhythms were less clear directly after anthropogenic food deprivation, in terms of the circadian rhythm in gape angle becoming significantly more apparent over the following days. Unlike mussels fed at midnight, those fed at midday displayed no significant change in gape angle from the hour before to the hour after they were fed, i.e. mussels given food at midday reacted to this food less than mussels fed at midnight. We suggest that independent of feeding time, laboratory mussels exposed to natural light and free from anthropogenic disturbance increase feeding activity at night because their circadian rhythms are strongly influenced by light levels. This study emphasises that the behaviour of animals in the laboratory and in the wild can be altered by anthropogenic disturbances such as vibrations caused by experimental setups and artificial illumination at night.     

Evidence of cannibalism and bentho-pelagic coupling within the life cycle of the mussel, Perna canaliculus

The feeding ecology of the green-lipped mussel, Perna canaliculus, was investigated within three intertidal mussel beds along Ninety Mile Beach, northern New Zealand, between August 2000 and March 2001. Adult mussels of different sizes (45-105 mm in shell length) were collected from the intertidal sites about 30 min after being submerged by the incoming tide for gut content analyses. Results of these analyses indicate that mussels consume a variety of phytoplankton, micro- and mesozooplankton, including mussel larvae and post-larvae. Cannibalism of juveniles of up to 620 μm was recorded for intertidal mussels, and conspecifics of up to 2.4 mm were found within the stomachs of additional mussels collected in August 2000 from a nearby subtidal site. For all three intertidal populations, mussel larvae and juveniles contribute about 70% of the food particle consumption during the spawning peak in August, while phytoplankton and other zooplankton constitute the majority of the food source (about 99%) in March, during gametogenesis. Larger intertidal mussels tended to have more food particles in their stomachs than smaller mussels within all three populations. Distinctive differences in food consumption among intertidal populations directly coincide with variations in total particulate matter (TPM), particulate organic matter (POM) and percent organic matter (OM) in the adjacent seawater.Separate experiments designed to test the feeding behavior of mussels feeding at different times during the incoming tide were conducted at one of the intertidal sites during August 2000 and March 2001. Results from these experiments indicate a marked shift in food consumption from bivalves to other mesozooplankton in August, and from phytoplankton to mesozooplankton in March. The observed combination of mussel predatory and grazing behavior over the incoming tide and through the year provides evidence for a strong food-web link between the benthic and pelagic life stages of this species. Furthermore, the high rate of cannibalism during some months of the year suggests that this source of food may significantly contribute to the energy budget of wild populations, with potential implications for evolutionary adaptive success.     

Life history traits and abundance can predict local colonisation and extinction rates of freshwater mussels

1. A critical need in conservation biology is to determine which species are most vulnerable to extinction. Freshwater mussels (Bivalvia: Unionacea) are one of the most imperilled faunal groups globally. Freshwater mussel larvae are ectoparasites on fish and depend on the movement of their hosts to maintain connectivity among local populations in a metapopulation. 2. I calculated local colonisation and extinction rates for 16 mussel species from 14 local populations in the Red River drainage of Oklahoma and Texas, U.S. I used general linear models and AIC comparisons to determine which mussel life history traits best predicted local colonisation and extinction rates. 3. Traits related to larval dispersal ability (host infection mode, whether a mussel species was a host generalist or specialist) were the best predictors of local colonisation. 4. Traits related to local population size (regional abundance, time spent brooding) were the best predictors of local extinction. The group of fish species used as hosts by mussels also predicted local extinction and was probably related to habitat fragmentation and host dispersal abilities. 5. Overall, local extinction rates exceeded local colonisation rates, indicating that local populations are becoming increasingly isolated and suffering an ‘extinction debt’. This study demonstrates that analysis of species traits can be used to predict local colonisation and extinction patterns and provide insight into the long‐term persistence of populations.     

Dynamic measurements of black oystercatcher (Haematopus bachmani) predation on mussels (Mytilus californianus)

Intertidal zone mussels can face threats from a variety of predatory species during high and low tides, and they must balance the threat of predation against other needs such as feeding and aerobic respiration. Black oystercatchers (Haematopus bachmani) on the Pacific coast of North America can depend on the mussel Mytilus californianus for a substantial portion of their diet. Observations suggest that oystercatchers tend to focus on mussels beginning to gape their valves during rising tides, following periods of aerial emersion. We present detailed, autonomous field measurements of the dynamics of three such predation events in the rocky intertidal zone. We measured accelerations of up to 4 g imposed on mussels, with handling times of 115–290 s required to open the shell and remove the majority of tissue. In each case a single oystercatcher attacked a mussel that had gaped the shell valves slightly wider than its neighbors as the rising tide began to splash the mussel bed, but no other obvious characteristic of the mussels, such as body temperature or orientation, could be linked to the oystercatcher's individual prey choice.     

Time budgets of Nuthatches Sitta europaea with supplementary food

Individual time budgets during winter and spring were studied in a population of Nuthatches Sitta europaea in south-central Sweden. In late winter birds spent about 90% of daytime foraging, males feeding somewhat less than females, possibly due to male dominance. In early spring, foraging time decreased to about 65% of daytime activity, and sex-related differences became more pronounced. Males then increased time spent resting and singing. Females were nestbuilding, spent less time resting and also devoted more time to foraging than did males. Prior to the observation periods, food density for part of the population was experimentally increased by supplying sunflower seed. In spite of this, time budgets in the experimental group were similar to those of the control group. It is possible that the intensity of foraging, rather than actual time devoted to it, was influenced by food density and that, as a result, birds in the experimental group were able to maintain a higher level of vigilance. Also, an increase in food supply may be less important to a hoarding bird such as the Nuthatch. However, the result remains that Nuthatch time budgets in winter and spring are not readily influenced by an increase in food density.     

Average annual growth and condition of mussels as a function of food source

Prior to 1986, before completion of a storm surge barrier, the average annual condition of mussels at culture plots in the Eastern Scheldt was better in the western part near the North Sea, than in the central part of the estuary. No such difference was observed in 1986 and 1987.To predict the effects of the barrier on mussel culture, the condition of the mussels, which is an index of growth rate, is analysed and correlated with food sources for the period 1981–84. In the western and central part of the Eastern Scheldt, mussel condition correlates strongly with average annual primary production, but not with chlorophyll-a concentrations. This suggests a direct link between primary production and the growth of mussels.In the western part of the estuary, the relation between condition and primary production has a less steep slope than in the central part. Import of food from the North Sea is suggested to act as an additional food source in the west of the Eastern Scheldt; hence the better condition values of the mussels, also in years with relatively low primary production.The storm surge barrier reduces the water exchange with the North Sea. Reduced import of food and consequently lower mussel condition are expected in the western part of the estuary. Preliminary data from 1987 confirm this prediction.     

Oystercatchers use colour preference to achieve longer-term optimality

The optimal diet model entails that foragers look beyond the individual prey encounter, to at least the level of intake rate across a bout of foraging, but optimization over a longer time remains controversial. In this paper, we show how oystercatchers increase their intake over the longer term using mussel colour as a cue. Wintering oystercatchers Haematopus ostralegus feed extensively on mussels Mytilus edulis in the estuaries of southern Britain. They show a marked preference for brown-shelled mussels over the commoner black-shelled morph, and we show that this enables them to maximize their rate of energy gain over a longer period than a single foraging bout. The brown and black mussels did not differ in ventral thickness and energy content, which are the main criteria for mussel selection and most important for short-term optimization. The brown mussels contained significantly less moisture, so by selecting them, oystercatchers could pack more mussel flesh into their limited oesophageal storage capacity. This enables them to increase their overall consumption during a feeding bout and increases their long-run energy gain rate, to an extent that is large enough to be significant for survival, especially during the short exposure of the mussel beds in winter.