Object Exploration of Garden and Sardinian Warblers Peaks in Spring

Seasonality of environments is a widespread phenomenon and many species show seasonal behavioural changes as an adaptation to this environmental variability. Exploratory behaviour, an important means to obtain information about the environment, varies between species depending on different costs and benefits of exploration under various environmental conditions or life styles. It can be expected that exploration also varies intraspecifically over the annual cycle according to seasonally changing costs and benefits of exploration due to seasonal environments and/or an organism's requirements. Captive garden and Sardinian warblers were confronted with five different and novel objects in their familiar aviary over the course of 1 yr. Both species showed a seasonal peak in object exploration in spring and low exploration values throughout the rest of the year indicating that knowledge about the environment is particularly important during the time of territory or nest‐site selection. Furthermore, the year‐round resident Sardinian warblers were more explorative than the migratory garden warblers. Residents have to be well informed about their environment and changes therein, whereas this is less important for migrants. This corroborates earlier findings in parrots and seems to be a general phenomenon consistent across taxa.     

Head Colour and Age Relate to Personality Traits in Gouldian Finches

Colour polymorphism is a widespread phenomenon in vertebrates and has often been linked with differences in behaviour such as aggression or boldness, behaviours that are often part of personality traits in monomorphic species. However, up to now, very few studies have looked whether colour morphs just differ in average behaviour or whether specific combinations of behaviours are favoured in relation to a particular morph therefore signalling personality traits. This was tested in the highly social and polymorphic Gouldian finch (Erythrura gouldiae). Tests were performed in pairs of a red‐headed and a black‐headed individual of same sex and size but different age. Capture rank in the aviary as well as latency to feed beside a novel object (neophobia) and to approach and touch a novel object on a perch (neophilia) were measured. For neophobia and neophilia, ranks of latencies calculated across all individuals were used and general and within‐pair comparisons were made. Neophobia and approach neophilia were consistent over time (2 mo), and approach neophilia was positively correlated with capture rank and neophobia indicating that the behaviours are part of personality traits. Moreover, black‐headed birds generally approached the novel object earlier than red‐headed birds. Similarly, within pairs, black‐headed birds were first to approach the novel object and were also captured first. Relative age (younger or older than the partner) was related to neophobia and tactile neophilia ranks; older birds reacted faster. The results indicate that colour morphs not only differ in average behaviour but that specific combinations of behaviours are related to colour morphs suggesting that head colours signal personality traits. Furthermore, depending on age individuals may have different roles in a social network.     

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.     

Agonistic behaviour,responses to a novel object and some aspects of maintenance behaviour in feral-strain and domestic chickens

Feral-strain fowl, bred from birds captured on North-West Island (24°S, 150°E), were compared with domestic chickens hatched and raised under similar conditions. Aspects of behaviour considered included agonistic behaviour and responses to unfamiliar environments and objects. There were many similarities between the feral and domestic fowl in behaviour, but there were also many differences between the strains.Feral cockerels showed higher levels of agonistic behaviour than domestic cockerels, under some conditions. There were differences between young feral and Leghorn-cross chickens in the “freezing” response to handling and placement in an unfamiliar cage, with the feral chickens responding more rapidly.Feral cockerels showed a greater initial avoidance of a novel object than did Black Australorp bantam cockerels, but after a short interval they spent more time near the novel object than the bantams.These results are discussed in relation to selection pressures under conditions of domestication, and on North-West Island.     

Relevance of individual and environmental drivers of movement of Golden Eagles

An animal's movement is expected to be governed by an interplay between goals determined by its internal state and energetic costs associated with navigating through the external environment. Understanding this ecological process is challenging when an animal moves in two dimensions and even more difficult for birds that move in a third dimension. To understand the dynamic interaction between the internal state of an animal and the variable external environment, we evaluated hypotheses explaining association of different covariates of movement and the trade-offs birds face as they make behavioural decisions in a fluctuating landscape. We used ~870 000 GPS telemetry data points collected from 68 Golden Eagles Aquila chrysaetos to test demographic, diel, topographic and meteorological hypotheses to determine (1) the probability that these birds would be in motion and (2), once in motion, their flight speed. A complex and sometimes interacting set of potential internal and external factors determined movement behaviour. There was good evidence that reproductive state, manifested as age, sex and seasonal effects, had a significant influence on the probability of being in motion and, to a lesser extent, on speed of motion. Likewise, movement responses to the external environment were often unexpectedly strong. These responses, to northness of slope, strength of orographic updraft and intensity of solar radiation, were regionally and temporally variable. In contrast to previous work showing the role of a single environmental factor in determining movement decisions, our analyses support the hypothesis that multiple factors simultaneously interact to influence animal movement. In particular they highlighted how movement is influenced by the interaction between the individual's internal reproductive state and the external environment, and that, of the environmental factors, topographic influences are often more relevant than meteorological influences in determining patterns of flight behaviour. Further disentangling of how these internal and externals states jointly affect movement will provide additional insights into the energetic costs of movement and benefits associated with achieving process-driven goals.     

Numerical and behavioural responses of migrant passerines to experimental manipulation of resident tits (Parus spp.): heterospecific attraction in northern breeding bird communites?

Summary We studied experimentally interspecific competition among foliage-gleaning passerine birds by manipulating the density of resident tits. In 1988 tit density was experimentally increased on three small islands in a central Finnish lake, and decreased on three other islands by tit removal. In order to avoid the effects of between-island differences in habitat quality, the role of the islands was reversed when the experiment was repeated in the following year. Censuses and observations on foraging and feeding behaviour were conducted to assess the numerical and behavioural responses of migrant conguilders (mainly chaffinches and willow warblers) with respect to the manipulated abundance of the tits. We also measured whether variation in food consumption of tits affected the frequency with which the migrants found food by calculating average intervals between successful prey captures, time lags to prey-capture and giving-up times. Our results indicate that interspecific competition is of minor importance in structuring breeding bird assemblages and species feeding ecologies on the study islands. No consistent difference in foraging or feeding niches of chaffinches and willow warblers was found between low and high tit density conditions. Niche overlap analysis showed no avoidance by chaffinches and willow warblers of the microhabitats which tits used. Tit abundance had no significant effect on feeding success or behaviour. Experimentally increased abundance of resident birds was associated with increased abundance of breeding migrants, however. This pattern was found not only in the foliage gleaning guild but also with all passerine birds, indicating that food was not an important contributor to this pattern. We elaborate a hypothesis suggesting heterospecific attraction in northern breeding bird assemblages. Habitat generalist migrants may use the presence of residents as an indicator of safe and/or productive breeding sites in northern unpredictable circumstances.     

Behavioural and endocrine fear responses in Japanese quail upon presentation of a novel object in the home cage

Most tests used to study fear in birds involve transferring them to a novel environment, which constitutes a bias in studies aiming at identifying the neural correlates of a specific fear-inducing situation. In order to investigate fear in birds with minimum interference by humans, behavioural and endocrine responses to the presentation of a novel object in the home cage were investigated in two lines of Japanese quail divergently selected for long or short duration of tonic immobility, a behavioural index of fear. Presentation of the novel object induced typical fear responses (avoidance of the object, increased pacing and increased plasma corticosterone levels) that were similar in the two lines of quail. Presentation of a novel object in the home cage thus appears to be a suitable stimulus to induce fear reactions in quail, with minimum interference from other motivational systems. The fact that quail of both lines reacted similarly in this test, while they are known to differ greatly in their behavioural responses to other fear-inducing tests, illustrates the multidimensional nature of fear.     

Exploratory behaviour and stressor hyper-responsiveness facilitate range expansion of an introduced songbird

Global anthropogenic changes are occurring at an unprecedented rate; one change, human-facilitated introduction of species outside their native range, has had significant ecological and economic impacts. Surprisingly, what traits facilitate range expansions post-introduction is relatively unknown. This information could help predict future expansions of introduced species as well as native species shifting their ranges as climate conditions change. Here, we asked whether specific behavioural and physiological traits were important in the ongoing expansion of house sparrows (Passer domesticus) across Kenya. We predicted that birds at the site of initial introduction (Mombasa, introduced approx. 1950) would behave and regulate corticosterone, a stress hormone, differently than birds at the range edge (Kakamega, approx. 885 km from Mombasa; colonized within the last 5 years). Specifically, we predicted greater exploratory behaviour and stronger corticosterone response to stressors in birds at the range edge, which may facilitate the identification, resolution and memory of stressors. Indeed, we found that distance from Mombasa (a proxy for population age) was a strong predictor of both exploratory behaviour and corticosterone release in response to restraint (but only while birds were breeding). These results suggest that certain behavioural and neuroendocrine traits may influence the ability of species to colonize novel habitats.     

SURVIVAL COST OF AN EARLY IMMUNE SOLICITING IN NATURE

If immune functions confer obvious benefits to hosts, life-history theory assumes that they also induce costs, leading to trade-offs between immunity and other fitness components. However, whether substantial fitness costs are associated with immune systems in the wild is debatable, as numerous factors may influence the costs and benefits associated with immune activation. Here, we explore the survival cost of immune deployment in postfledging birds. We injected Eurasian collared dove nestlings ( Streptopelia decaocto ) with antigens from Escherichia coli , and examined whether this immune challenge affected survival after fledging. To assess survival, birds were fitted with radiotags and the fate of each individual was monitored regularly. Our results show that mimicking a bacterial infection in nestlings lowered their survival prospects after fledging, in comparison to controls. The main identified cause of mortality (by examination of dead birds) was presumed to be predation. This study provides experimental evidence that immune activation may entail dramatic survival costs in a free-ranging vertebrate, and emphasizes the potential role that environmental factors such as predation may play in this interaction.     

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.     

The importance of pattern similarity between Müllerian mimics in predator avoidance learning

Müllerian mimicry, where unpalatable prey share common warning patterns, has long fascinated evolutionary biologists. It is commonly assumed that Müllerian mimics benefit by sharing the costs of predator education, thus reducing per capita mortality, although there has been no direct test of this assumption. Here, we specifically measure the selection pressure exerted by avian predators on unpalatable prey with different degrees of visual similarity in their warning patterns. Using wild-caught birds foraging on novel patterned prey in the laboratory, we unexpectedly found that pattern similarity did not increase the speed of avoidance learning, and even dissimilar mimics shared the education of naive predators. This was a consistent finding across two different densities of unpalatable prey, although mortalities were lower at the higher density as expected. Interestingly, the mortalities of Müllerian mimics were affected by pattern similarity in the predicted way by the end of our experiment, although the result was not quite significant. This suggests that the benefits to Müllerian mimics may emerge only later in the learning process, and that predator experience of the patterns may affect the degree to which pattern similarity is important. This highlights the need to measure the behaviour of real predators if we are to understand fully the evolution of mimicry systems.     

Effects of Group Size on Approach to Novel Objects in Ravens (Corvus corax)

Neophobia may constrain explorative behaviour, learning and innovation, while social context may facilitate approach to novel objects and acceptance of novel food. We examined the effects of neophobia on the exploration of novel objects in relation to social context in ravens (Corvus corax). Ravens are suitable subjects for studying effects of social context, as they are highly neophobic scavengers that recruit conspecifics to food. We tested two groups of six and 11 hand‐raised birds in three conditions: single‐bird, in dyadic combinations and sibling sub‐groups of three to six birds. Contrary to expectation, individuals of both groups were quicker to approach novel objects when tested alone than when tested with conspecifics. However, they spent more time close to and manipulating the novel objects in the social conditions (dyadic and group) than when being alone. We discuss the possibility that the higher latencies of dyads and groups to approach novel objects may reflect a ‘negotiation’ process in a ‘war of attrition’ between the individuals over risk‐taking.     

Influence of a novel object in the home range of the cockroach,Blattella germanica

We investigated whether exploratory behaviour of the cockroach, Blattella germanica (L.) (Dictyoptera: Blattellidae), was modified by the introduction of a novel object, such as a bait station, into their familiar home range. In particular, we measured the attractiveness of boxes, as novel objects, in relation to their complexity and to their contents. The presence of a novel object induced an increase of exploratory behaviour on and inside the object and its environs. Time spent around the object depended on the complexity of the object (closed box or with holes). The more complex the object, the longer cockroaches spent exploring it. When the object contained food, its attractiveness was greatly increased and it affected cockroaches from further away.     

Once bitten twice shy: long‐term behavioural changes caused by trapping experience in willow warblers Phylloscopus trochilus

In dangerous situations, animals learn from experience to anticipate risks during similar subsequent occasions. Mist netting may represent just such a dangerous occasion and may serve as a tool to get insight into whether and how memorizing and recognition of the circumstances that form the prelude to a recapture can alter the animal's behaviour after capture, as well as it may affect the subsequent recapture rates. We used a playback lure to study the capture latency in willow warblers Phylloscopus trochilus, both without any and with a previous capture experience (either in the same year or in a previous one). We found increased response latencies to the playback lure for experienced males, compared to naive males. Naive males responded faster in the presence of a mist net than without a mist net; while experienced males tended to increase their response latencies. Individuals with between‐year capture experience showed exactly the same response pattern as individuals with a capture experience within the same year. These findings suggest that the birds may recall the capture event even a year later. It is very likely that the playback lure, with or without the net's presence, contributed significantly to the apparent risk of detection. We believe our study reveals that long‐lasting modifications of behaviour induced by capture and handling deserve further attention, as they might have a serious impact on the value of behavioural and ecological data, which is either based upon observing captured individuals and/or upon the recapture rates.     

Habitat suitability and the constraints of migration in New World warblers

Identifying the factors that influence geographic range limits can illustrate the various ecological, physiological, and evolutionary constraints imposed on a species. The range limits of migratory birds are particularly challenging to study as they occur in disjunct regions at different times of the year, which can impose different constraints. Travel between breeding and wintering regions poses a significant energetic and navigational challenge to birds, although it is not clear how these movements influence breeding dispersal and range expansion. Here I ask whether the possible costs of migration limit the breeding ranges of wood warblers, a group of birds with an extensive history of ecological and evolutionary studies. I used occurrence records for multiple wood warbler species, breeding primarily in the boreal forest of North America, to generate environmental niche models. I tested for over‐prediction of habitat suitability into the western boreal forest, where most these species do not have occurrence records but where there is presumably suitable habitat. I found that some of these vagile taxa, primarily found east of the Rocky Mountains, also have predicted habitat suitability that extends into the north and west, where they have little to no occurrence records. I discuss several possible explanations for this discordance. In particular, the patterns are consistent with the costs of a long‐distance migration limiting the benefits of range expansion, as migration may become too onerous as the distance between breeding and wintering areas increases. These results speak to the process of niche filling more generally and call attention to an under‐appreciated explanation for why migratory species may not fully occupy their fundamental niche.     

Delayed behavioral effects of postnatal exposure to corticosterone in the zebra finch (Taeniopygia guttata)

Early developmental conditions can significantly influence the growth and survival of many animal species. We studied the consequences of exposure to corticosterone (CORT), a stress hormone, during the nestling stage on two behavioral traits (neophobia, social dominance) measured when the birds had reached independence. Nestling zebra finches (Taeniopygia guttata) were exposed twice daily to exogenous CORT via oral administration for a 12-day period up until fledging. Experimental CORT administration depressed nestling growth rates, confirming results previously obtained in this species. Our data on neophobic behavior revealed a significant interaction between sex and treatment, with CORT-dosed males showing reduced latencies to approach a novel object, while there was little effect of corticosterone treatment on female neophobia. There was no significant effect of age (30 or 50 days), however, there was a non-significant trend towards an interaction between treatment and age, with neophobia increasing with age in the CORT-dosed birds, but decreasing in controls. At 50 days of age previous exposure to corticosterone resulted in reduced success in competitions for a non-food-based resource (a perch) in both sexes. There were no effects of brood size on any behavioral traits measured here, but this may be due to the small range in brood size used. Our results show that elevated levels of stress hormones during postnatal development can have significant effects on important behavioral traits, i.e., neophobia and dominance. Moreover, they confirm the importance of rearing conditions in shaping adult phenotypes.     

Urbanization and its effects on personality traits: a result of microevolution or phenotypic plasticity?

Human‐altered environmental conditions affect many species at the global scale. An extreme form of anthropogenic alteration is the existence and rapid increase of urban areas. A key question, then, is how species cope with urbanization. It has been suggested that rural and urban conspecifics show differences in behaviour and personality. However, (i) a generalization of this phenomenon has never been made; and (ii) it is still unclear whether differences in personality traits between rural and urban conspecifics are the result of phenotypic plasticity or of intrinsic differences. In a literature review, we show that behavioural differences between rural and urban conspecifics are common and taxonomically widespread among animals, suggesting a significant ecological impact of urbanization on animal behaviour. In order to gain insight into the mechanisms leading to behavioural differences in urban individuals, we hand‐raised and kept European blackbirds (Turdus merula) from a rural and a nearby urban area under common‐garden conditions. Using these birds, we investigated individual variation in two behavioural responses to the presence of novel objects: approach to an object in a familiar area (here defined as neophilia), and avoidance of an object in a familiar foraging context (defined as neophobia). Neophilic and neophobic behaviours were mildly correlated and repeatable even across a time period of one year, indicating stable individual behavioural strategies. Blackbirds from the urban population were more neophobic and seasonally less neophilic than blackbirds from the nearby rural area. These intrinsic differences in personality traits are likely the result of microevolutionary changes, although we cannot fully exclude early developmental influences.     

Hamilton's rule and the causes of social evolution

Hamilton''s rule is a central theorem of inclusive fitness (kin selection) theory and predicts that social behaviour evolves under specific combinations of relatedness, benefit and cost. This review provides evidence for Hamilton''s rule by presenting novel syntheses of results from two kinds of study in diverse taxa, including cooperatively breeding birds and mammals and eusocial insects. These are, first, studies that empirically parametrize Hamilton''s rule in natural populations and, second, comparative phylogenetic analyses of the genetic, life-history and ecological correlates of sociality. Studies parametrizing Hamilton''s rule are not rare and demonstrate quantitatively that (i) altruism (net loss of direct fitness) occurs even when sociality is facultative, (ii) in most cases, altruism is under positive selection via indirect fitness benefits that exceed direct fitness costs and (iii) social behaviour commonly generates indirect benefits by enhancing the productivity or survivorship of kin. Comparative phylogenetic analyses show that cooperative breeding and eusociality are promoted by (i) high relatedness and monogamy and, potentially, by (ii) life-history factors facilitating family structure and high benefits of helping and (iii) ecological factors generating low costs of social behaviour. Overall, the focal studies strongly confirm the predictions of Hamilton''s rule regarding conditions for social evolution and their causes.     

Roost site selection by Little Owls Athene noctua in relation to environmental conditions and life‐history stages

Roost site selection is a state‐dependent process, affected by the individual's costs and benefits of roosting at a specific site in the available environment. Costs and benefits of different roost sites vary in relation to intrinsic factors and environmental conditions. Thus, the cost–benefit functions of roost sites are expected to differ between seasons and life‐history stages, resulting in adjustments in roost site selection. Studying roost site selection throughout the year therefore provides information about year‐round habitat requirements at different life‐history stages. However, little is known about the roosting behaviour of birds. Here, the roost site selection of Little Owls Athene noctua was studied by repeated daytime location of 24 adult and 75 juvenile radiotagged individuals from July to November. Little Owls preferred sheltered roost sites such as tree cavities with multiple entrances. They increasingly used sheltered sites from summer to winter and preferentially used sheltered roost sites with low ambient temperatures. Juveniles used significantly less sheltered sites during dispersal than before and afterwards, and used less sheltered sites than adults within their home‐range. The survival probability of birds roosting frequently at exposed sites was reduced. Roost site selection is probably driven by the two mechanisms of predator avoidance and thermoregulation, and the costs of natal dispersal may include increased predation threat and higher energy expenditure for thermoregulation. We suggest that adequate roost sites, such as multi‐entrance tree cavities, are an important habitat requirement for Little Owls and that habitat quality can be affected by manipulating their availability.     

Hunt or hide: How insularity and urbanization affect foraging decisions in lizards

Foraging decisions should reflect a balance between costs and benefits of alternative strategies. Predation risk and resource availability in the environment may be crucial in deciding how cautious individuals should behave during foraging. These costs and benefits will vary in time and context, meaning that animals should be able to adjust their foraging behaviour to new or altered environments. Studying how animals do this is essential to understand their survival in these environments. In this study, we investigated the effect of both insularity and urbanization on risk‐taking and neophobia during foraging in the Dalmatian wall lizard (Podarcis melisellensis). Small islets tend to have both a lower number of predators and less resources. Therefore, islet populations were expected to show more risk‐taking behaviour and less neophobia in a foraging context. Previous studies on behaviour of urban lizards have yielded inconsistent results, but due to a lack of both predators and arthropod prey in urban habitats, we expected urban lizards to also take more risks and behave less neophobic. We sampled several inhabited and uninhabited locations on Vis (Croatia) and surrounding islets. Risk‐taking behaviour was tested by measuring the latency of lizards to feed in the presence of a predator model, and neophobia by measuring the latency to feed in the presence of a novel object. We found that islet lizards do indeed take more risks and were less vigilant, but not less neophobic. Urban and rural lizards did not differ in any of these behaviours, which is in sharp contrast with previous work on mammals and birds. The behavioural differences between islet and island lizards were novel, but not unexpected findings and are in line with the theory of “island tameness”. The effect of urbanization on the behaviour of animals seems to be more complex and might vary among taxa.