Experience with adults shapes multisensory representation of social familiarity in the brain of a songbird

Social animals learn to perceive their social environment, and their social skills and preferences are thought to emerge from greater exposure to and hence familiarity with some social signals rather than others. Familiarity appears to be tightly linked to multisensory integration. The ability to differentiate and categorize familiar and unfamiliar individuals and to build a multisensory representation of known individuals emerges from successive social interactions, in particular with adult, experienced models. In different species, adults have been shown to shape the social behavior of young by promoting selective attention to multisensory cues. The question of what representation of known conspecifics adult-deprived animals may build therefore arises. Here we show that starlings raised with no experience with adults fail to develop a multisensory representation of familiar and unfamiliar starlings. Electrophysiological recordings of neuronal activity throughout the primary auditory area of these birds, while they were exposed to audio-only or audiovisual familiar and unfamiliar cues, showed that visual stimuli did, as in wild-caught starlings, modulate auditory responses but that, unlike what was observed in wild-caught birds, this modulation was not influenced by familiarity. Thus, adult-deprived starlings seem to fail to discriminate between familiar and unfamiliar individuals. This suggests that adults may shape multisensory representation of known individuals in the brain, possibly by focusing the young's attention on relevant, multisensory cues. Multisensory stimulation by experienced, adult models may thus be ubiquitously important for the development of social skills (and of the neural properties underlying such skills) in a variety of species.     

Behavioral, physiological, and endocrine responses of starlings to acute increases in density

One potential stressor to vertebrates both in the wild and in captivity is the presence of numerous individuals in a confined space. To examine the effects of increased conspecific density in birds, we simultaneously measured cardiac, behavioral, and endocrine responses of European starlings (Sturnus vulgaris) to acute crowding. A cage containing a resident bird was outfitted with a trap door that allowed for the introduction of intruder birds (one, three, or five birds) without human interference. The resident bird was implanted with a subcutaneous heart rate (HR) transmitter, behavior was videotaped through a two-way mirror, and blood samples were taken at the end of each treatment to determine plasma corticosterone (CORT) concentrations. Resident starlings significantly increased both general activity and aggressive behaviors while decreasing preening following the initiation of elevated conspecific density. Intruder starlings increased feeding, drinking, and aggressive pecking rates, but postintrusion feeding rates decreased as intruder number increased. Preening decreased in both residents and intruders following the intrusion. HR increased in the resident starlings at the time of intruder introduction, with an increase in the magnitude of this response directly correlating with increasing intruder number. The CORT response to increased density was dependent on social role (resident or intruder), since increasing density did not alter CORT levels in resident birds, but resulted in elevated CORT 30 min following the five-intruder introduction in the intruder birds. Together, these data suggest that increased conspecific density is a significant acute stressor in starlings which is capable of inducing aggression in both residents and intruders. Furthermore, it elicits different responses from different physiological and behavioral systems, and behavioral responses such as feeding and general activity may be density-dependent. The data specifically illustrate that cardiac and behavioral activation can be independent of CORT release, and the CORT response of starlings to increased conspecific density is dependent on social role and degree of the increase in density.     

Mirror-mediated responses of California scrub jays (Aphelocoma californica) during a caching task and the mark test

Mirror self-recognition, as an index of self-awareness, has been proposed as a precursor for more complex social cognitive abilities, such as prosocial reasoning and cooperative decision-making. Indeed, evidence for mirror self-recognition has been shown for animals possessing complex social cognitive abilities such as great apes, dolphins, elephants and corvids. California scrub jays (Aphelocoma californica) have provided strong evidence that non-human animals are capable of mental state attribution. For instance, scrub jays are reported to use their experience stealing the food of others to infer that other birds may similarly intend to steal from them. If a concept of “self” is required for such complex social cognitive abilities, then scrub jays might be expected to show mirror self-recognition. Thus, we examined whether California scrub jays are capable of mirror self-recognition using two experimental contexts: a caching task and the mark test. During the caching task, we compared the extent to which scrub jays protected their food after caching alone, in the presence of a conspecific and in the presence of a mirror. The birds did not engage in more cache protection behaviours with a mirror present than when caching alone, suggesting scrub jays may have recognized their reflection and so did not expect cache theft. Alternative explanations for this behaviour are also discussed. During the mark test, the scrub jays were surreptitiously marked with a red or plumage-coloured control sticker. The scrub jays showed no evidence of mirror self-recognition during the mark test, as the birds did not preferentially attempt to remove the red mark in the presence of a mirror. Together, the results provide mixed evidence of the mirror self-recognition abilities of California scrub jays. We highlight the need to develop alternative approaches for evaluating mirror self-recognition in non-human animals to better understand its relationship with complex social cognition.     

Fear and exploration in European starlings (Sturnus vulgaris): a comparison of hand-reared and wild-caught birds

The revision of EU legislation will ban the use of wild-caught animals in scientific procedures. This change is partially predicated on the assumption that captive-rearing produces animals with reduced fearfulness. Previously, we have shown that hand-reared starlings (Sturnus vulgaris) indeed exhibit reduced fear of humans compared to wild-caught conspecifics. Here, we asked whether this reduction in fear in hand-reared birds is limited to fear of humans or extends more generally to fear of novel environments and novel objects. Comparing 6-8 month old birds hand-reared in the lab with age-matched birds caught from the wild as fledged juveniles a minimum of 1 month previously, we examined the birds' initial reactions in a novel environment (a small cage) and found that wild-caught starlings were faster to initiate movement compared to the hand-reared birds. We interpret this difference as evidence for greater escape motivation in the wild-caught birds. In contrast, we found no differences between hand-reared and wild-caught birds when tested in novel object tests assumed to measure neophobia and exploratory behaviour. Moreover, we found no correlations between individual bird's responses in the different tests, supporting the idea that these measure different traits (e.g. fear and exploration). In summary, our data show that developmental origin affects one measure of response to novelty in young starlings, indicative of a difference in either fear or coping style in a stressful situation. Our data contribute to a growing literature demonstrating effects of early-life experience on later behaviour in a range of species. However, since we did not find consistent evidence for reduced fearfulness in hand-reared birds, we remain agnostic about the welfare benefits of hand-rearing as a method for sourcing wild birds for behavioural and physiological research.     

Behavioural and hormonal effects of social isolation and neophobia in a gregarious bird species, the European starling (Sturnus vulgaris)

Separating gregarious individuals from their group members often results in behavioural and physiological changes, like increased levels of corticosterone. Testosterone and corticosterone, in particular, have been implicated in the response of mammals to novelty. Data in birds are, however, rare. The presence or absence of group members may also influence an individual's response to novel stimuli. We assessed the behaviour and hormonal response of European starlings (Sturnus vulgaris) to a novel object in two different situations and seasons: each starling was tested when separated and when in contact with its group members in May/June (breeding season) and again in September/October (non-breeding season). Starlings are gregarious throughout the year, but as foraging flocks are small during the breeding season and large during the non-breeding season, we assumed that non-breeding starlings would be more affected by social isolation. Overall, starlings had higher levels of corticosterone, lost more body mass, and were more active when they were separated from their group. Isolated individuals, however, did not show a greater neophobic response than individuals in the presence of their group members in either season. Circulating levels of testosterone and corticosterone were higher after a test with novel object than after a test with only the familiar feeding dish in both sexes and seasons. However, control tests for handling effects confirmed only the increase in testosterone. Our study shows that social isolation is stressful for unrelated and unpaired members of a wild flocking bird species and demonstrates that novelty can lead to a rise in testosterone in birds.     

Are Female Starlings Able to Recognize the Scent of Their Offspring?

Background

Although there is growing evidence that birds may have individual chemical profiles that can function in several social contexts, offspring recognition based on olfactory cues has never been explored. This ability should be more likely evolved in colonial birds and/or species suffering brood parasitism, in which the risk of being engaged in costly misdirected parental care is high.

Methodology/Principal Findings

We performed a choice experiment to examine whether females of the spotless starling, Sturnus unicolor, a species that is colonial, and where a fraction of the population is exposed to intraspecific brood parasitism, can discriminate between the scent of their offspring and that of unrelated nestlings. We also explored whether the development of the uropygial gland secretion may play a role in such olfactory discrimination by performing the choice experiments to females rearing nestlings of two different ages, that is, without and with developed uropygial glands. Results showed that female starlings did not preferentially choose the scent of their offspring, independently of whether the gland of nestlings was developed or not.

Conclusions/Significance

Our results suggest that female starlings do not have or do not show the ability to distinguish their offspring based on olfaction, at least up to 12–14 days of nestling age. Further research is needed to examine whether odour-based discrimination may function when fledgling starlings leave the nest and the risk of costly misidentification is likely to increase.     

Safety first: terrestrial predators drive selection of highly specific nesting sites in colonial‐breeding birds

Nesting is a critical yet hazardous life stage for many birds. For colonial‐breeding birds, the conspicuousness of the colony to predators suggests immense pressure to select optimal colonial nesting sites. But what drives selection of those sites? As with solitary nesting birds, reducing access by predators may be the single most important factor. If so, knowledge of the predators involved and the attributes of different potential colony sites can allow us to predict the features that make a site especially safe. We examined the attributes of trees used by breeding colonies of metallic starlings Aplonis metallica in tropical Australia, and experimentally tested if those attributes prevented nest access by predatory snakes. Our surveys confirmed that tree choice by starling nesting colonies is highly non‐random, with all colonies located in tall trees in rainforest clearings, with no low branches and smooth bark. Experimental tests demonstrated that the climbing ability of predatory snakes depends upon bark rugosity, and that colony access by snakes depends on tree attributes such as bark rugosity and canopy connectivity. Our study confirms that colonial‐nesting starlings select colony sites that provide a safe refuge from predation. Intense predation pressure may have driven the evolution of stringent breeding habitat criteria in many other species of colonial‐breeding birds.     

Use and husbandry of captive European starlings (Sturnus vulgaris) in scientific research: a review of current practice

We reviewed the use of captive European starlings (Sturnus vulgaris) in scientific research published between 2000 and 2004. We estimated the numbers of birds used and documented their origin and the range of husbandry regimes employed with the aim of comparing current practice with the new European guidelines for husbandry of laboratory animals. Over the five-year period, 106 primary articles report the use of an estimated total of 2490 captive starlings. The majority of birds were caught from the wild either as adults or fledglings, and only 3% were hand-reared from chicks. There was considerable variation in husbandry. In the majority of cases, standards fell below those currently recommended as best practice in the UK and cited in new European guidelines. The median volume of home cages employed was 0.42 m3 (0.13-5.1 m3, interquartile range), whereas current recommendations suggest a minimum of 1.0 m3 for a singly-housed bird. The median volume of space allowed per bird was 0.13 m3/bird (0.08-1.05 m3/bird, Q1-Q3), whereas current recommendations suggest a minimum of 0.33 m3/bird. Only 27% of the articles mentioned providing any form of environmental enrichment for birds in their home cages. We recommend that more research be conducted into the welfare of starlings to inform legislation and guidelines, and thus maximize the welfare of captive animals.     

Efficacy of European starling control to reduce <Emphasis Type="Italic">Salmonella enterica</Emphasis> contamination in a concentrated animal feeding operation in the Texas panhandle

Background  

European starlings (Sturnus vulgaris) are an invasive bird species known to cause damage to plant and animal agriculture. New evidence suggests starlings may also contribute to the maintenance and spread of diseases within livestock facilities. Identifying and mitigating the risk pathways that contribute to disease in livestock is necessary to reduce production losses and contamination of human food products. To better understand the impact starlings have on disease transmission to cattle we assessed the efficacy of starling control as a tool to reduce Salmonella enterica within a concentrated animal feeding operation. We matched a large facility, slated for operational control using DRC-1339 (3-chloro-4-methylaniline hydrochloride, also 3-chloro p-toluidine hydrochloride, 3-chloro-4-methylaniline), with a comparable reference facility that was not controlling birds. In both facilities, we sampled cattle feed, cattle water and cattle feces for S. enterica before and after starling control operations.     

Experimental analysis of the social value of flocking by starlings (Sturnus vulgaris) in relation to predation and foraging

In groups of ten, indidual starlings, Sturnus vulgaris, spent significantly less time in surveillance than did individuals in smaller groups and responded more quickly than single birds to a flying model hawk. Captive starlings in flocks reduce their individual surveillance efforts, but their combined efforts still enable them to be more effective than single birds in the detection of predators. Foraging behaviour of flocks was observed by placing single starlings with groups of tricoloured blackbirds, Agelaius tricolor; the starlings reduced the time they devoted to surveillance at the same rate as if they were with other starlings.     

Preferences and apparent digestibilities of sugars by fruit damaging birds in Japan

Gray starlings Sturnus cineraceus, azure-winged magpies Cyanopica cyana and brown-eared bulbuls Hypsipetes amaurotis are among the main bird pests in commercial fruit orchards in central Japan. Recently Brugger & Nelms (1991) suggested that developing high-sucrose fruit cultivars could reduce crop damage, because some pest birds lack the enzyme sucrase and can develop an aversion to sucrose. Preferences for, and digestibilities of, the monosaccharides glucose and fructose and the disaccharide sucrose by these pests species were therefore studied to assess whether this idea would be applicable in central Japan. Gray starlings and brown-eared bulbuls were able to detect glucose, fructose, a mixture of glucose and fructose, and sucrose at a concentration of 12% w/v. Azure-winged magpies also detected glucose and fructose, but failed to detect sucrose at the same concentration. In pairwise preference trials gray starlings and azure-winged magpies selected the monosaccharides over sucrose, but brown-eared bulbuls did not. To estimate the digestibility of the sugars the apparent assimilated mass coefficient, AMC*, was calculated for each species eating each sugar by measuring intake and faecal output. Monosaccharides had mean AMC*s of 0.77, 0.96 and 0.92 when consumed by gray starlings, azure-winged magpies and brown-earned bulbuls respectively. AMC* values for sucrose were 0.82 and 0.49 for brown-eared bulbuls and azure-winged magpies respectively, but gray starlings were shown to be unable to digest sucrose. As AMC* values varied from 0.75 to 0.97, consumption rates of sugars increased as digestibility decreased. Although increasing sucrose contents of commercial fruits may deter sucrase-deficient birds such as gray starlings from depredating fruits, it may also lead to increased crop damage by species such as the azure-winged magpie and brown-eared bulbul which may have to consume more of the less digestible fruit in order to meet their energy requirements.     

Avian influenza A virus susceptibility,infection, transmission,and antibody kinetics in European starlings

Avian influenza A viruses (IAVs) pose risks to public, agricultural, and wildlife health. Bridge hosts are spillover hosts that share habitat with both maintenance hosts (e.g., mallards) and target hosts (e.g., poultry). We conducted a comprehensive assessment of European starlings (Sturnus vulgaris), a common visitor to both urban and agricultural environments, to assess whether this species might act as a potential maintenance or bridge host for IAVs. First, we experimentally inoculated starlings with a wild bird IAV to investigate susceptibility and replication kinetics. Next, we evaluated whether IAV might spill over to starlings from sharing resources with a widespread IAV reservoir host. We accomplished this using a specially designed transmission cage to simulate natural environmental transmission by exposing starlings to water shared with IAV-infected mallards (Anas platyrhynchos). We then conducted a contact study to assess intraspecies transmission between starlings. In the initial experimental infection study, all inoculated starlings shed viral RNA and seroconverted. All starlings in the transmission study became infected and shed RNA at similar levels. All but one of these birds seroconverted, but detectable antibodies were relatively transient, falling to negative levels in a majority of birds by 59 days post contact. None of the contact starlings in the intraspecies transmission experiment became infected. In summary, we demonstrated that starlings may have the potential to act as IAV bridge hosts if they share water with IAV-infected waterfowl. However, starlings are unlikely to act as maintenance hosts due to limited, if any, intraspecies transmission. In addition, starlings have a relatively brief antibody response which should be considered when interpreting serology from field samples. Further study is needed to evaluate the potential for transmission from starlings to poultry, a possibility enhanced by starling’s behavioral trait of forming very large flocks which can descend on poultry facilities when natural resources are scarce.     

Behavioral and physiological responses of wild-caught European starlings (Sturnus vulgaris) to a minor, rapid change in ambient temperature

Free-ranging animals continuously adjust to changes in their environment. The stress response, typified by increases in heart rate and glucocorticoids, is an important physiological response regulating these changes. This study investigated heart rate, corticosterone and behavioral responses of European starlings (Sturnus vulgaris) to a rapid 30min decrease in temperature using an air-conditioning unit. Ten wild-caught birds were divided into pairs and exposed to four different trials. Three trials were controls: undisturbed birds; exposing birds to only the noise of the air-conditioning unit; and exposing the birds to 20°C airflow. For the experimental trial birds were exposed to 12°C air, leading to a rapid but modest 3°C drop in ambient temperature inside the birdcages. Heart rate and behavior were recorded before and during trials, while blood samples were collected before and after each trial for corticosterone measurements. Cooling, but none of the control conditions, induced an increase in heart rate and corticosterone. Additionally, cooling led to an increase in perch hopping and feather ruffling. We conclude that minor changes in temperature can elicit a stress response in European starlings, which suggests that this may be an important mechanism by which animals cope with minor rapid environmental changes.     

Independent effects of song quality and experience with photostimulation on expression of the immediate,early gene ZENK (EGR‐1) in the auditory telencephalon of female European starlings

Age influences behavioral decisions such as reproductive timing and effort. In photoperiodic species, such age effects may be mediated, in part, by the individual's age‐accrued experience with photostimulation. In female European starlings (Sturnus vulgaris) that do not differ in age, experimental manipulation of photostimulation experience (photoexperience) affects hypothalamic, pituitary, and gonadal activity associated with reproductive development. Does photoexperience also affect activity in forebrain regions involved in processing a social cue, the song of males, which can influence mate choice and reproductive timing in females? Female starlings prefer long songs over short songs in a mate‐choice context, and, like that in other songbird species, their auditory telencephalon plays a major role in processing these signals. We manipulated the photoexperience of female starlings, photostimulated them, briefly exposed them to either long or short songs, and quantified the expression of the immediate‐early gene ZENK (EGR‐1) in the caudomedial nidopallium as a measure of activity in the auditory telencephalon. Using an information theoretic approach, we found higher ZENK immunoreactivity in females with prior photostimulation experience than in females experiencing photostimulation for the first time. We also found that long songs elicited greater ZENK immunoreactivity than short songs did. We did not find an effect of the interaction between photoexperience and song length, suggesting that photoexperience does not affect forebrain ZENK‐responsiveness to song quality. Thus, photoexperience affects activity in an area of the forebrain that processes social signals, an effect that we hypothesize mediates, in part, the effects of age on reproductive decisions in photoperiodic songbirds. © 2009 Wiley Periodicals, Inc. Develop Neurobiol 2009     

Immediate early gene activity in song control nuclei and brain areas regulating motivation relates positively to singing behavior during, but not outside of, a breeding context

In some species, such as songbirds, much is known about how the brain regulates vocal learning, production, and perception. What remains a mystery is what regulates the motivation to communicate. European starlings (Sturnus vulgaris) sing throughout most of the year, but the social and environmental factors that motivate singing behavior differ seasonally. Male song is highly sexually motivated during, but not outside of, the breeding season. Brain areas outside the song control system, such as the medial preoptic nucleus (POM) and ventral tegmental area (VTA), have been implicated in regulating sexually motivated behaviors in birds, including song. The present study was designed to explore whether these regions, as well as three song control nuclei [area X, the high vocal center (HVC), and the robust nucleus of the arcopallium (RA)], might be involved differentially in song produced within compared to outside of a breeding context. We recorded the behavioral responses of breeding and nonbreeding condition male starlings to the introduction of a female conspecific. Males did not show context-dependent differences in the overall amount of song sung. However, immunocytochemistry for the protein product of the immediate early gene cFOS revealed a positive linear relationship between the total amount of songs sung and number of cFOS-labeled cells in POM, VTA, HVC, and RA for birds singing during, but not outside of, a breeding context. These results suggest that these regions differentially regulate male song production depending on reproductive context. Overall the data support the hypothesis that the POM and VTA interact with the song control system, specifically HVC and RA, to regulate sexually motivated vocal communication in songbirds.     

Do European starlings prefer light environments containing UV?

Many captive birds are kept in artificial lighting that is typically deficient in ultraviolet (UV) wavelengths. Most birds can perceive the range of light that humans see but also have an additional retinal cone type that is tuned to UV wavelengths. Consequently, artificial lighting may be detrimental as it might limit the functionality of their vision. We examined the preferences of European starlings, Sturnus vulgaris, for various artificial light environments. In our first experiment, groups of starlings showed a preference for environments that contained UV (UV+) over those where UV wavelengths had been removed (UV−). This preference was not affected by the sex of the individuals within the group or, as shown in a later experiment, by whether the birds had been previously housed in UV+ or UV− conditions. In contrast, individual starlings showed no preference for UV+ over UV− environments, although the power of our test was low. In a subsequent experiment, starling groups preferred the higher of two light intensities that were presented; however, equalizing the overall quantal flux between UV+ and UV− extinguished any preference for UV+ over UV−. The group preference for UV+ conditions in the first experiment may therefore have resulted from a preference for brighter conditions rather than a specific preference for UV. However, equalizing the quantal flux may not equalize perceived brightness, because it is not known how birds' visual systems weight input from each cone type. We conclude that, for nonbreeding, group-housed captive starlings, there is no positive evidence of a preference for the presence of UV as a specific wavelength. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Are the low protein requirements of nectarivorous birds the consequence of their sugary and watery diet? A test with an omnivore

Nectar-feeding birds have remarkably low nitrogen requirements. These may be due either to adaptation to a low-protein diet or simply to feeding on a fluid diet that minimizes metabolic fecal nitrogen losses. We measured minimal nitrogen requirements (MNR) and total endogenous nitrogen loss (TENL) in the omnivorous European starling Sturnus vulgaris, fed on an artificial nectar-like fluid diet of varying concentrations of sugar and protein. The MNR and TENL of the birds were similar and even slightly higher than allometrically expected values for birds of the starlings' mass (140% and 103%, respectively). This suggests that the low measured nitrogen requirements of nectar-feeding birds are not simply the result of their sugary and watery diets but a physiological adaptation to the low nitrogen input. We also measured the effect of water and protein intake on the nitrogenous waste form in the excreta and ureteral urine in European starlings. Neither high water intake nor low protein intake increased the fraction of nitrogen excreted as ammonia. Ammonia was excreted at consistently low levels by the starlings, and its concentration was significantly higher in ureteral urine than in excreta. We hypothesize that ureteral ammonia was reabsorbed in the lower intestine, indicating a postrenal modification of the urine.     

Social learning in birds and its role in shaping a foraging niche

We briefly review the literature on social learning in birds, concluding that strong evidence exists mainly for predator recognition, song, mate choice and foraging. The mechanism of local enhancement may be more important than imitation for birds learning to forage, but the former mechanism may be sufficient for faithful transmission depending on the ecological circumstances. To date, most insights have been gained from birds in captivity. We present a study of social learning of foraging in two passerine birds in the wild, where we cross-fostered eggs between nests of blue tits, Cyanistes caeruleus and great tits, Parus major. Early learning causes a shift in the foraging sites used by the tits in the direction of the foster species. The shift in foraging niches was consistent across seasons, as showed by an analysis of prey items, and the effect lasted for life. The fact that young birds learn from their foster parents, and use this experience later when subsequently feeding their own offspring, suggests that foraging behaviour can be culturally transmitted over generations in the wild. It may therefore have both ecological and evolutionary consequences, some of which are discussed.     

Captive European starlings (Sturnus vulgaris) in breeding condition show an increased cardiovascular stress response to intruders

European starlings (Sturnus vulgaris) alter their physiology and behavior between seasons, becoming territorial during the spring/summer and flocking during the fall/winter. We used captive male starlings in breeding (photostimulated to 18L : 6D) and nonbreeding (11L : 13D) conditions to determine whether changing physiology and behavior alters their reaction to crowding. One or five intruders entered a resident's cage without human disturbance. A subcutaneous heart rate transmitter recorded cardiovascular output in residents. Corticosterone and testosterone were measured in plasma samples taken before and after the intrusion. While corticosterone concentrations did not change, heart rate changed significantly, indicating that these responses can be regulated independently. Long-day birds showed a significantly elevated heart rate response to the single-bird intrusion compared to short-day birds. Whereas five intruders elicited an identical peak response in both groups, long-day birds also demonstrated an equivalent response to one intruder. In addition, one intruder induced longer elevation in heart rate for long-day birds. Male starlings in breeding condition, therefore, demonstrate an increased sensitivity to additional conspecifics. This seasonal shift in response suggests that a higher tolerance for intrusion (i.e., considering a nearby starling as less stressful) may facilitate flocking behavior, while a lower tolerance may aid in territoriality.     

Searching in patches by European starlings,Sturnus vulgaris

Despite the importance of within-patch search for predicting optimal patch-leaving strategy, little experimental effort has been devoted to the study of this foraging behaviour. In addition, predators' strategic responses to variability in features like the within-patch distribution of prey can have important consequences for their dietary decisions. We therefore analyse the search paths of European starlings, Sturnus vulgaris, foraging individually in artificial patches that vary in the spatial distribution of buried prey in an outdoor aviary. The results demonstrate that the birds are searching differently depending on the order in which the experimental patch-types are experienced. We speculate that where the spatial predictability of prey is initially high the birds adopt a fixed search rule that results in area-concentrated search once a prey item is found, and performs well for both distributions encountered. However, where the predictability is initially low a more flexible strategy is adopted that results in increased area-concentrated search with experience of a patch type, independent of the actual within-patch distribution of prey. These findings suggest that starlings can use distinct strategies for different prey types, but they are classifying these types on subjective criteria that are difficult to predict from a priori reasoning.