Mussel dropping by Carrion and Hooded crows: biomechanical and energetic considerations

Dropping live mussels (Mytilus sp.) onto hard substrata by Carrion Crows (Corvus corone) and Hooded Crows (Corvus cornix) to access their flesh is a commonly observed behavior from late summer to spring in the United Kingdom and Ireland. Despite previous studies, several aspects of prey‐dropping behavior remained incompletely understood. From September 2008 to January 2010, we determined the heights of drops, likelihood of shell breakage from drops at different heights, effect of mussel size on breakability, energetic costs of flying to drop heights, and the energetic costs of transporting mussels from mussel beds to dropping sites. We studied Carrion Crows on the Isle of Cumbrae, Scotland, and Hooded Crows in Cork Harbor, Ireland. Initial experiments were carried out with mussels to determine breakability in relation to size and drop height, and to estimate mussel energy content. Sizes of mussel shells at Hooded Crow dropping sites were compared with those of live mussels from source mussel beds. Adult Carrion Crows (N = 10) dropped mussels from a mean height of 4.7 m, and adult Hooded Crows (N = 21) from 4.8 m. These heights were close to the minimum (4–4.8 m; determined experimentally) required to break all mussel shells on the first drop. Dropping mussels from the minimum height that guarantees breakage reduces handling time and, by minimizing the size of the resulting debris field, likely reduces the risk of kleptoparasitism. Juvenile Hooded Crows (N = 13) dropped mussels onto suboptimal substrates (gravelly mud) from variable heights (mean = 6.1 m) with a low success rate (0% on first drop). This inefficiency could reflect either inexperience or exclusion from prime hard‐substrate dropping sites by adults. Foraging Hooded Crows selected larger mussels, dropping no mussels <32‐mm shell length. Energetic calculations indicate that a Hooded Crow lifting a medium‐sized mussel (55‐mm shell length) to a height of 5 m incurs a cost of only 0.3% of energy assimilated from that mussel, whereas travel to and from a mussel bed 200 m away costs 5.8% of that energy. These results suggest that choice of mussel dropping height by crows is determined by shell breakability rather than the cost of flying up to the dropping height.     

Avian prey-dropping behavior. I. The effects of prey characteristics and prey loss

Numerous species of birds break hard-shelled prey items by droppingthem from a height. This intriguing prey-extraction method providesan excellent opportunity for studying foraging behavior becausea single, easily measurable quantity—height of drop—maybe influenced by a wide variety of identifiable characteristicsof the prey (e.g., breakability, weight) and social environment(e.g., alone or in the presence of kleptoparasites). Using adynamic, state variable modeling approach, this paper presentsthe first theoretical framework for avian prey-dropping systemsthat incorporates the diversity of prey characteristics andsocial situations. The model yielded a series of qualitativepredictions about prey-dropping behavior that can be testedreadily in any prey-dropping system. In particular, the results indicatedthat quantitative and qualitative differences in item breakability andpotential kleptoparasitism should have a significant effecton the height and pattern of prey dropping.     

Context-dependent ‘safekeeping’ of foraging tools in New Caledonian crows

Several animal species use tools for foraging, such as sticks to extract embedded arthropods and honey, or stones to crack open nuts and eggs. While providing access to nutritious foods, these behaviours may incur significant costs, such as the time and energy spent searching for, manufacturing and transporting tools. These costs can be reduced by re-using tools, keeping them safe when not needed. We experimentally investigated what New Caledonian crows do with their tools between successive prey extractions, and whether they express tool ‘safekeeping’ behaviours more often when the costs (foraging at height), or likelihood (handling of demanding prey), of tool loss are high. Birds generally took care of their tools (84% of 176 prey extractions, nine subjects), either trapping them underfoot (74%) or storing them in holes (26%)—behaviours we also observed in the wild (19 cases, four subjects). Moreover, tool-handling behaviour was context-dependent, with subjects: keeping their tools safe significantly more often when foraging at height; and storing tools significantly more often in holes when extracting more demanding prey (under these conditions, foot-trapping proved challenging). In arboreal environments, safekeeping can prevent costly tool losses, removing a potentially important constraint on the evolution of habitual and complex tool behaviour.     

Effects of different dropping intensities on fascicle and tendinous tissue behavior during stretch-shortening cycle exercise.

This study examined whether the elasticity of the tendinous tissues plays an important role in human locomotion by improving the power output and efficiency of skeletal muscle. Ten subjects performed one-leg drop jumps (DJ) from different dropping heights with a constant rebound height. The fascicle length of the vastus lateralis muscle was measured by using real-time ultrasonography during DJ. In the braking phase of the DJ, fascicle lengthening decreased and the tendinous tissue lengthening increased with increased dropping intensity. In the subsequent push-off phase, the shortening of tendinous tissues increased with higher dropping intensity. The averaged electromyographic activities of the preactivation and braking phases increased and those of the push-off phase decreased as the drop height was increased. With higher dropping height but constant submaximal rebound jump, the stretched tendinous tissue length increased with less stretched fascicle during the braking phase. In the subsequent push-off phase, the recoil of tendinous tissues became greater. These results suggest that the increased prestretch intensity has considerable influence on the process of storage and subsequent recoil of the elastic energy during the stretch-shortening cycle action.     

Substrate optimization in nut cracking by capuchin monkeys (Cebus apella)

We conducted an experiment to examine the effect of substrate on the nut-cracking behavior of a group of semicaptive capuchin monkeys. We wanted to determine whether tufted capuchin monkeys were selective in choosing the substrate on which they pound nuts, and whether the choice of substrate affected the outcome. Eight adult females and eight juveniles were provided with nuts in the outdoor facility. We found that 1) all adult females and three young capuchins succeeded in cracking nuts; 2) they preferred the hardest substrates (concrete and stone); 3) there is a link between the substrate and the amount of time needed to crack a nut; 4) most young capuchins used various substrates, some of which were inadequate, in a haphazard manner; and 5) there are different forms of nut cracking. We conclude that adult capuchins choose the hardest substrates, and that these substrates support efficient cracking.     

Do kleptoparasites reduce their own foraging effort in order to detect kleptoparasitic opportunities?: An empirical test of a key assumption of kleptoparasitic models

Determining if, or when, individuals trade off time spent personally feeding against time spent monitoring others for kleptoparasitism opportunities is essential to an understanding of the evolution of scrounging and usurpation behaviours. We provide a first field test of whether kleptoparasites reduce their personal foraging effort in situations where the frequency and rewards of kleptoparasitism increase. We provided experimental food patches for wild European blackbirds that varied in the distribution of prey and that had a potentially high rate of kleptoparasitism within pairs of blackbirds feeding in them. Although individuals differed in their rate of kleptoparasitism, they did not vary in the size of the reward that they gained from kleptoparasitism. As prey became more clumped, kleptoparasitism rate and its reward per incident increased on average. There was, however, no evidence that individuals that were kleptoparasitising more quickly and/or at a higher frequency had lower personal foraging effort. In contrast, foraging effort increased in both birds compared to when they were foraging alone, independent of dominance, kleptoparasitic opportunity or reward. Our evidence suggests that in some circumstances a kleptoparasite can detect kleptoparasitic opportunities without compromising its own personal foraging rate.     

The effect of kleptoparasite and host numbers on the risk of food‐stealing in an avian assemblage

Kleptoparasitism involves the theft of resources such as food items from one individual by another. Such food‐stealing behaviour can have important consequences for birds, in terms of individual fitness and population sizes. In order to understand avian host–kleptoparasite interactions, studies are needed which identify the factors which modulate the risk of kleptoparasitism. In temperate European intertidal areas, Eurasian oystercatchers Haematopus ostralegus feed primarily on bivalve molluscs, which may be stolen by kleptoparasitic species such as carrion crows Corvus corone and herring gulls Larus argentatus. In this study we combined overwinter foraging observations of oystercatchers and their kleptoparasites on the Exe Estuary, UK, with statistical modelling to identify the factors that influence the likelihood of successful food stealing behaviour occurring. Across the winter, 16.4% of oystercatcher foraging attempts ended in successful kleptoparasitism; the risk of theft was lowest in February (10.8%) and highest in December (36.3%). Using an information theoretic approach to compare multiple logistic regression models we present evidence that the outcome of host foraging attempts varied with the number of kleptoparasites per host within the foraging patch for two out of five individual months, and for all months grouped. Successful, kleptoparasitism was more likely to occur when the total number of all kleptoparasites per host was greater. Across the entire winter study period, oystercatcher foraging attempts that resulted in kleptoparasitism were associated with a mean number of kleptoparasites per host that was more than double that for foraging attempts that ended in the oystercatcher successfully consuming the mussel. Conversely, the stage of the tidal cycle within the estuary did not affect the outcome of oystercatcher foraging attempts. Our study provides evidence that bird numbers influence the risk of kleptoparasitism within avian assemblages.     

Seedlessness and Parthenocarpy inPistacia veraL. (Anacardiaceae): Temporal Changes in Patterns of Vascular Transport to Ovules

Pistacia vera‘Kerman’ (pistachio nut) typicallyproduces high numbers of seedless or blank fruits. Patternsof vascular transport into fruits and ovules were studied over3 years by following the movement of disodium fluorescein solutionfrom cut branches into developing fruitlets. Early in the season,vascular conductivity is intact through to the chalazal endof the ovule. Soon afterwards, the percentage of ovules withvascular conductivity through to the chalaza declines, and ina variable fraction of fruits, movement of the fluorochromesolution becomes blocked either at the placenta or in the funiculus.Six to 9 weeks after anthesis there is blockage in 90 (1 year)to 100% (2 years) of fruits. Subsequently, vascular conductivityresumes in 83.3% (3 year mean) of ovules, a percentage thatcorrelates well with the mean percentage of seeded nuts at harvest(77.5%). Ovules from fruits with dysfunctional vascular conductionearly in the season are smaller than those with fully functionalvascular tissue. At the time conductivity declines, a high percentageof those ovules with blocked vascular movement lack endospermand appear to be unfertilized; none of the ovules that retainfull vascular flow lack endosperm. Pollination using gamma-irradiatedpollen (⁶⁰Co, 1.0 kGy) led to a nearly three-fold increase inthe production of blank nuts. The results indicate that fluoresceintransport may be a valuable tool to predict the fate of ovules,and are consistent with the hypothesis that parthenocarpic fruitset may be an important factor in blank nut production in pistachio.Copyright1999 Annals of Botany Company Pistachio,Pistacia veraL., fluorescein, seed set, seedlessness, parthenocarpy, blanking, ovule, funiculus, chalaza, embryo.     

Auditory prey location in a pause--travel predator: search height, search time, and attack range of Tengmalm's owls (Aegolius funereus)

We studied the nocturnal hunting and diurnal roosting behaviorof 17 radio-equipped Tengmalm's owls (Aegolius funereus), 12males and 5 females, in coniferous forest during their nestingseason. The owls perched lower when hunting than when roosting,probably because hunting perches were selected to minimize thepredator-prey distance or to obtain unobstructed access to theground-dwelling small mammal prey, whereas roosting percheswere selected to minimize the probability of being detectedby an avian predator. There was no difference between perchingheights associated with giving up and prey attack, nor werethere any differences between perching heights, perching times,and attack distances associated with successful and unsuccessfulattacks. There were no sexual differences in perching heightduring hunting or roosting. However, giving-up times tendedto be longer for females than for their mates, which is expectedbecause females are larger than males, and the relative costof flight increases with body mass. The instantaneous attackrate was independent of perching time. The owls gave up theirperches at a constant rate and independently of the amount oftime already spent on the perch in an exponentially decayingpattern. The owls perched longer, however, before launchingan attack than before giving up, probably in order to observedetected prey until the right moment for an attack. Attack distancewas independent of both perching height and perching time. Perchingtime was inversely related to perching height, which fits thetheoretical expectation that the search area will decrease withincreasing height in birds that locate prey auditorily.     

The evolutionary ecology of nut dispersal

A variety of nut-producing plants have mutualistic seed-dispersal interactions with animals (rodents and corvids) that scatter hoard their nuts in the soil. The goals of this review are to summarize the widespread horticultural, botanical, and ecological literature pertaining to nut dispersal inJuglans, Carya, Quercus, Fagus, Castanae, Castanopsis, Lithocarpus, Corylus, Aesculus, andPrunus; to examine the evolutionary histories of these mutualistic interactions; and to identify the traits of nut-bearing plants and nut-dispersing rodents and jays that influence the success of the mutualism. These interactions appear to have originated as early as the Paleocene, about 60 million years ago. Most nuts appear to have evolved from ancestors with wind-dispersed seeds, but the ancestral form of dispersal in almonds (Prunus spp.) was by frugivorous animals that ingested fruit. Nut-producing species have evolved a number of traits that facilitate nut dispersal by certain rodents and corvids while serving to exclude other animals that act as parasites of the mutualism. Nuts are nutritious food sources, often with high levels of lipids or proteins and a caloric value ranging from 5.7 to 153.5 kJ per propagule, 10–1000 times greater than most wind-dispersed seeds. These traits make nuts highly attractive food items for dispersers and nut predators. The course of nut development tends to reduce losses of nuts to insects, microbes, and nondispersing animals, but despite these measures predispersal and postdispersal nut mortality is generally high. Chemical defenses (e.g., tannins) in the cotyledons or the husk surrounding the nut discourage some nut predators. Masting of nuts (periodic, synchronous production of large nut crops) appears to reduce losses to insects and to increase the number of nuts dispersed by animals, and it may increase cross-pollination. Scatter hoarding by rodents and corvids removes nuts from other sources of nut predation, moves nuts away from source trees where density-dependent mortality is high (sometimes to habitats or microhabitats that favor seedling establishment), and buries nuts in the soil (which reduces rates of predation and helps to maintain nut viability). The large nutrient reserves of nuts not only attract animal dispersers but also permit seedlings to establish a large photosynthetic surface or extensive root system, making them especially competitive in low-light environments (e.g., deciduous forest) and semi-arid environments (e.g., dry mountains, Mediterranean climates). The most important postestablishment causes of seedling failure are drought, insufficient light, browsing by vertebrate herbivores, and competition with forbs and grasses. Because of the nutritional qualities of nuts and the synchronous production of large nut crops by a species throughout a region, nut trees can have pervasive impacts on other members of ecological communities. Nut-bearing trees have undergone dramatic changes in distribution during the last 16,000 years, following the glacial retreat from northern North America and Europe, and the current dispersers of nuts (i.e., squirrels, jays, and their relatives) appear to have been responsible for these movements.     

Studies on Mechano-Perception in Characean Cells: Development of a Monitoring Apparatus

An apparatus to monitor the electrical signal generated in Characells upon mechanical stimulation was developed. An internodalcell was stimulated by dropping a glass tubing with the intensityof the stimulus being controlled by changing either the weightof the glass tubing or the height from which it was dropped.Upon stimulation, a receptor potential was generated with theamplitude being dependent on the stimulus intensity. When thereceptor potential reached a threshold value, an action potentialwas generated. The receptor potential and action potential werecharacterized. The usefulness of this apparatus for analyzingreceptor potentials is discussed. (Received January 29, 1996; Accepted April 16, 1996)     

Evolutionarily stable kleptoparasitism: consequences of different prey types

We present two elaborations of the model of Broom and Ruxtonthat found evolutionarily stable kleptoparasitic strategiesfor foragers. These elaborations relax the assumption that thedistribution of times required to handle discovered food itemsis exponential. These changes increase the complexity of themodel but represent a significant improvement in biologicalrealism. In one elaboration, handling takes a fixed interval,t_h, at the end of which the whole value of the food item isobtained. We liken this to peeling then consuming a small orange.The other elaboration also assumes that handling takes a fixedinterval, t_h, but this time the reward from the food item isextracted continuously throughout the handling period. We likenthis to eating an apple. Both models predict that increasingfood density, the ease with which food items can be discovered,or the length of aggressive contests all act to make kleptoparasitismless common. The difference between the evolutionarily stablestrategy solutions of the apple and orange models provides aclear prediction of our theory. When prey items require handlingbefore yielding a lump sum at the end, then kleptoparasiticattacks will be focused on prey items near the end of theirhandling period. However, if prey items yield reward continuouslyduring handling, then attacks should be biased toward newlydiscovered food items. Another key difference between the modelpredictions is that kleptoparasitism increases with foragerdensity in the apple model, but decreases in the orange model.     

Foraging Decision Rules and Prey Species Preferences of Northwestern Crows (Corvus caurinus)

Categorization of similar prey types and the application of decision rules by dietary generalists can enhance the efficiency of foraging decisions and facilitate the inclusion of novel prey types in the diet. While considerable research attention has been directed toward investigation of these concepts in invertebrates, few have assessed categorization and decision rules used by generalist vertebrate predators. In this study, we experimentally investigated decision rules and prey preferences of northwestern crows (Corvus caurinus) feeding on littleneck clams (Tapes philippinarum) and whelks (Nucella lamellosa). We presented crows with three species‐size combinations: small clams (2.0–2.9 cm length) paired with large whelks (4.0–4.9 cm), small clams paired with medium whelks (3.0–3.9 cm), and large clams (4.0–4.9 cm) with large whelks. Profitability estimates based on observations of crows feeding on these prey species indicated that clams were always the more energetically profitable option; however, in prey choice trials crows consistently selected the heavier prey species, regardless of differences in profitability. These results show that crows apply a general decision rule according to which they select heavier prey items when feeding on hard‐shelled prey requiring similar handling techniques, and that while such decision rules may approximate optimal choices they may not always follow predictions based solely on prey profitability. We discuss these results in the context of behavioural flexibility of generalist predators, and predicting impacts of intertidal avian predators on prey populations.     

Experimental field test of spatial variation in rodent predation of nuts relative to distance and seed density

The spatial context in which seed predation occurs may modify the spatial structure of recruitment generated by seed dispersal. The Janzen–Connell (J-C) model predicts that granivores will exert greater pressure on the parent plant or at those sites where the density of dispersed seeds is higher. We have investigated how the probability of post-dispersal survival of Juglans australis varies with nut density across a hierarchy of spatial scales. We experimentally evaluated the survival of 3,120 nuts at three spatial scales: meso-scale (≤1.5 ha), as forest sites with two densities of fruiting J. australis individuals; intermediate scale (<0.2 ha), as individual trees with two experimental crop sizes; small scale (<0.1 m²), as microsites with two factors (number of nuts and distance from source). Nut removal coincided with seed predation, a condition that allowed us to test the density-dependent seed predation hypothesis. We found that the probability of nut survival was greater at forest sites with higher J. australis density. Nut survival was not affected by nut density in the seed shadow of individual specimens: at sites where J. australis density was greater, the proportion of surviving nuts did not differ between microsites located at different distances from the parent plant, but it was greater at microsites with greater initial nut density. Nut survival depended on the scale at which rodents responded to nut density, being negatively density dependent at the meso-scale and spatially random at intermediate and small scales. At the meso-scale, excess nut supply increased the probability of nut survival, which is in agreement with a model of granivore satiation near the seed source. Rodent satiation at the meso-scale may favour maintenance of sites with high J. australis density, where individual trees may have greater probabilities of passing their genes onto the next stage of the dispersal cycle.     

Dispersal of sweet pignut hickory in a year of low fruit production,and the influence of predation by a curculionid beetle

Summary The rate at which fallen hickory nuts are removed from beneath the parent tree, and the effect on this rate of the seed predatorConotrachelus affinis, was studied in an oak-hickory forest in southeastern Michigan, USA, during a year in which few nuts were produced. The trees responded toConotrachelus, which destroyed half the nut crop, by aborting inviable nuts during the summer. The seed dispersers, mostly gray squirrels, removed fallen nuts rapidly, showing the ability to distinguish viable nuts and remove them preferentially. The number of nuts removed in a week varies directly with the number available, and removal rate increases when many viable nuts are falling. The death of most seeds before dispersal, and the squirrels' efficiency at foraging on nuts and recovering them after burial, imply that successful hickory reproduction takes place only in years of heavy nut production.     

The diet of coastal breeding hooded crows corvus corone comix

Three methods of analysis were used to determine the diet of territorial hooded crows at Lough Hyne Marine Reserve, Co Cork, Ireland The regular collection of prey Items from these sites at Lough Hyne was integrated with pellet and stomach analysis to determine diet Intertidal organisms occurred in over 80% of pellets and 43% of stomachs and occupied over 77% of the total wet weight of foods identified in pellets All prey items recovered from drop sites originated from the intertidal habitat, involved either large-sized species or larger individuals of smaller-sized species, and were only dropped during October to February Twenty-five intertidal species were identified but only a few of these species contributed to the bulk of the diet Hooded crows were shown to consume a wide range of intertidal species throughout the year, though the species composition in the diet was seasonally influenced Depletion and weight loss of intertidal molluscs through the winter was shown to have a minimal effect on selection suggesting that prey switching was driven by the birds nutritional requirements     

The role of chromatic and achromatic signals for fruit detection by birds

Fruit color changes during ripening are typically viewed asan adaptation to increase signal efficacy to seed dispersers.Plants can increase signal efficacy by enhancing chromatic (wavelengthrelated) and/or achromatic (intensity related) contrasts betweenfruit and background. To assess how these contrasts determinethe detectability of fruit signals, we conducted 2 experimentswith free-flying crows (Corvus ossifragus) under seminaturalconditions in a 2025 m² aviary. Crows searched first for artificialred and black fruits and detected red fruits from a larger distance.Because artificial red fruits had higher chromatic and lowerachromatic contrasts against foliage than artificial black fruits,crows apparently prioritized chromatic contrasts. Thus, thecommon change in fruit color from red to black during ripeningdoes not increase signal efficacy to crows. In a second trial,crows searched for UV-reflecting and black blueberries (Vaccinummyrtillus) against backgrounds of foliage and sand. Againstfoliage, UV-reflecting berries had higher chromatic and achromaticcontrasts than black berries, and crows detected them from alarger distance. Against sand, UV-reflecting berries had lowachromatic contrasts and black berries low chromatic contrasts.Crows detected both fruit types equally, suggesting that theyused chromatic contrasts to detect UV-reflecting berries andachromatic contrasts to detect black berries. Birds prioritizedchromatic contrasts when searching for artificial red fruitsin foliage but not when searching for blueberries on sand. Wesuggest that the relative importance of chromatic and achromaticcontrasts is contingent on the chromatic and achromatic varianceof the background. Models of signal perception can be improvedby incorporating background-specific effects.     

The use of territorial gardening versus kleptoparasitism by a subtropical reef fish (Kyphosus cornelii) is influenced by territory defendability

Many territorial herbivorous animals increase the productivityor quality of the prey community on their territory. However,this gardening tactic may be vulnerable to kleptoparasitismunless gardeners can exclude others from the patch. We investigatedthe influence of territory defendability on the decision ofwestern buffalo bream (Kyphosus cornelii) to defend an algalgarden or to leave their territories and invade those of conspecifics.Fish monitored while away from their territories did kleptoparasitizethe gardens of conspecifics. We manipulated the structural complexityof territories through the addition of short or tall simulatedvegetation. Increasing the height of simulated vegetation onthe territory decreased the defendability of territories butincreased their safety for fish; territories with tall vegetationwere more easily invaded by conspecifics, but fish returnedmore rapidly to these territories after being disturbed. Ina paired experiment, fish spent more time away from their territories(roving) when tall vegetation was added than when short vegetationwas added. However, this effect was influenced by depth, witha greater difference between treatments on deeper territories,and by body size, with a greater difference for larger fish.These results support the prediction that territory-holdingindividuals faced with a trade-off between defending their owngarden or kleptoparasitizing from others adjust their use ofthese tactics based on the defendability of resources.     

Evidence that tufted puffins Fratercula cirrhata use colony overflights to reduce kleptoparasitism risk

Predation, foraging and mating costs are critical factors shaping life histories. Among colonial seabirds, colony overflights may enhance foraging or mating success, or diminish the risk of predation and kleptoparasitism. The latter possibility is difficult to test because low predation or kleptoparasitism rates could be due either to low danger or to effective counter-tactics by prey. Tufted puffins Fratercula cirrhata breeding at a large colony in British Columbia, Canada, deliver several loads of fish each day to their nestlings and are targets for kleptoparasitism by glaucous-winged gulls Larus glaucescens . In the present study, we documented the ecological conditions under which overflights occurred in order to assess when overflights were made and to statistically isolate the effect of overflights on kleptoparasitism risk at this site. Load-carrying puffins engaged in overflights under ecological conditions associated with relatively high rates of kleptoparasitism. Further, when ecological factors determining risk were statistically controlled, overflights were correlated with marginally lower chances of kleptoparasitism than when the risk factors were ignored. The results suggest that breeding puffins at this site use overflights for kleptoparasite avoidance. This tactic is used sparingly, suggesting it is costly. Costs of overflight behaviour might contribute to the impact of kleptoparasitism on the breeding success of tufted puffins.     

Kleptoparasitism in gulls Laridae at an urban and a coastal foraging environment: an assessment of ecological predictors

Capsule: Kleptoparasitism in gulls occurred at a greater rate at an urban compared with a coastal site. Population density and prey size predicted the rate of kleptoparasitism at the urban site.

Aims: To investigate and assess the ecological variables associated with kleptoparasitism among gulls at urban and rural sites.

Methods: Field observations were conducted at Brancaster (coastal rural) and Billingsgate Market (urban) to examine differences in the rate of kleptoparasitism in mixed-species flocks of gulls. Four key variables (prey size, population density, season and species) were assessed as predictors of kleptoparasitism.

Results: Generalized linear models revealed significant effects on kleptoparasitism rate of site, population density and prey size, and two-way interactions between these main terms. Population density and prey size differed significantly between sites, but population density appeared to predict the rate of kleptoparasitism.

Conclusion: Kleptoparasitism may well aid invasion and increase the range of environments a gull can tolerate by helping them meet their energy needs in novel environments where normal foraging behaviours are difficult to implement.