Predator Detection is Limited in Microhabitats with High Light Intensity: An Experiment with Brown‐Headed Cowbirds

Variations in ambient light conditions across different microhabitats can modify the detectability of predators and prey. Prey have been shown to be more visible in sunlit than in shaded patches, leading to higher predation risk and more investment in vigilance (predation risk hypothesis). Additionally, prey have been hypothesized to take longer to detect predators in sunlit compared to shaded patches because of the excess of sunlight causing glare effects (disability glare hypothesis). We tested the predictions of these two non‐mutually exclusive hypotheses in a seminatural experiment with brown‐headed cowbirds by measuring vigilance behavior and detection of a ground predator in patches under the shade of vegetation and in the open. Light intensity and achromatic contrast were higher in the sunlit patches, which could enhance glare effects, but chromatic contrast was higher in the shaded patches. Brown‐headed cowbirds took longer to show alert reactions to and flee from a ground predator in sunlit compared to shaded patches. However, the two parameters associated with perceived predation risk (vigilance prior to the predator exposure and time to resume foraging after the attack) did not differ between sunlit and shaded patches. Our findings support to a greater extent the disability glare hypothesis than the predation risk hypothesis. Overall, ambient light conditions can affect two critical components of behavioral predator–prey interactions in terrestrial habitats: detection of and escape from predators. The effects of disability glare are expected to be more pronounced in bird species with wider visual fields or without sun‐shading structures; however, species may compensate through various behaviors (e.g. avoidance of sunlit patches and changes in head orientation).     

Does sun glare increase antipredator behaviour in prey?

As the sun gradually lowers over the horizon, prey species with more sun in their eyes should have more difficulty in visually monitoring their surroundings for threats and thus experience a higher predation risk. In a unique setting, I could examine changes in antipredator behaviour in a prey species, the semipalmated sandpiper Calidris pusilla, facing attacks by peregrine falcons Falco peregrinus, which originated from the general direction of the lowering sun. I predicted gradual changes in antipredator behaviour as sun glare becomes more problematic later in the day. As the day progressed, sandpipers occurred in sparser groups when the sun glared but not when clouds obscured the sun, suggesting that fewer individuals engaged in risky foraging. Pecking rate and foraging success decreased later in the day when the sun glared but not otherwise implying an increase in vigilance at the expense of foraging. When more sun hit their eyes, sandpipers also moved faster suggesting increased skittishness. The sun glare effect might be relevant to any species foraging in open areas not only when the sun sets but also when it rises especially if predators can target prey species at these vulnerable times. The temporal gradient in predation risk that the sun glare effect creates might thus apply broadly and have important consequences for antipredator vigilance, foraging efficiency, and habitat use.     

Light environment and the impacts of foliage quality on herbivorous insect attack and bird predation

Theory predicts that variation in plant traits will modify both the direct interactions between plants and herbivores and the indirect impacts of predators of those herbivores. Light has strong effects on leaf quality, so the impacts of herbivores and predators may differ between plants grown in sun and shade. However, past experiments have often been unable to separate the effects of light environment on plant traits and herbivory from direct effects on herbivores and predators. We first manipulated light availability in an open habitat using a shade cloth pre-treatment to produce oak saplings with different leaf qualities. Leaves on plants exposed to high light were thicker and tougher and had lower nitrogen and water contents, and higher carbon and phenolic contents than leaves on plants under a shade cloth. Then, in the main experiment, we moved all plants to a common shade environment where bird predators were excluded in a factorial design. We measured insect herbivore abundance and leaf damage. Herbivores were significantly more abundant and caused greater leaf damage on sun trees, although these leaf characteristics are usually associated with low-quality food. Bird exclusion did not change herbivore abundance but did increase leaf damage. Contrary to our predictions, the effects of birds did not differ between trees grown in sun and shade conditions. Thus, differences in effects of predators on herbivores and plants between light habitats, when observed, might be due to variation in predator abundance and not bottom-up effects of host plant quality.     

Pigeons in the sun: Thermal constraints of eumelanic plumage in the rock pigeon (Columba livia)

In wild vertebrates, several species exhibit eumelanic color polymorphism with the coexistence of dark and light morphs. The maintenance of such polymorphism suggests the existence of a selective balance between the morphs and a large body of literature has reported the costs and benefits of darker plumage coloration in birds. Among them, it has been suggested that melanin and dark plumage could entail high energetic costs especially under hot and sunny climates. However, to my knowledge, the thermal constraints of sun exposure have rarely been studied in polymorphic species. Here, we tested the impact of eumelanic plumage coloration on plumage and body temperatures, and evaporative cooling behavior in the polymorphic rock pigeon (Columbia livia). We experimentally exposed light and dark pigeons to direct sun radiation for 1 h while a few birds were maintained in the shade as controls. We found that sun exposure was associated with increased plumage temperature, and this effect was greater for darker pigeons. In addition, we found that sun exposure was also associated with higher cloacal temperature but for dark pigeons only. Finally, light and dark pigeons were more likely to show cooling evaporative behavior when exposed to sun and as their cloacal temperature increases. Altogether, these results suggest that darker pigeons may have a lower ability to cope with heat and solar radiations and that dark plumage can be associated with thermal costs in this polymorphic species.     

Thermal constraints on foraging in adult european starlings

Summary The operative temperature of the environment was estimated for starlings using hollow, unheated taxidermic mounts. On average, adults foraging in full sun were characterized by shorter foraging bouts than those adults foraging in full shade. Simultaneous observations of air temperature, operative temperature, and the foraging duration of adults indicated that air temperature was a poor predictor of the maximum length of a foraging bout. The operative temperature of the environment was not correlated to the maximum and mean length of foraging bouts for temperatures below 31.5°C, but was negatively related to maximum and mean foraging duration for values above 31.5°C. I also found that foraging adults experiencing high thermal loads (T _e31.5°C) were less likely to return to the nest with food. These results raise the possibility that adults may be limited in their utilization of foraging sites due to an inability to cope with heat load, and that much of the loafing behavior observed for breeding birds may be behavior to avoid heat stress. The implication for seasonal variation of clutch size is discussed.     

Does sunlight enhance the effectiveness of avian preening for ectoparasite control?

Preening is a bird's first line of defense against harmful ectoparasites. Ectoparasites, in turn, have evolved adaptations for avoiding preening such as hardened exoskeletons and escape behavior. Earlier work suggests that some groups of ectoparasites, such as feather lice, leave hiding places in feathers that are exposed to direct sunlight, making them more vulnerable to preening. It is, therefore, conceivable that birds may choose to preen in direct sunlight, assuming it improves the effectiveness of preening. Using mourning doves and their feather lice, we tested 2 related hypotheses; (1) that birds with access to direct sunlight preen more often than birds in shade, and (2) that birds with access to direct sunlight are more effective at controlling their ectoparasites than birds in shade. To test these hypotheses, we conducted an experiment in which we manipulated both sunlight and preening ability. Our results provided no support for either hypothesis, i.e., birds given the opportunity to preen in direct sunlight did not preen significantly more often, or more effectively, than did birds in shade. Thus, the efficiency of preening for ectoparasite control appears to be independent of light intensity, at least in the case of mourning doves and their feather lice.     

Response to Predation Risk in Urban and Rural House Sparrows

Habitat urbanization may change the density of predators, and it is often assumed that such changes lead to altered predation risk for urban populations of their prey. Although it is difficult to study predation hazard directly, behavior responses of prey species may be informative in inferring such habitat differences. In this study, we compared the risk‐taking behavior of urban and rural house sparrows (Passer domesticus) after simulated attacks by two of their important predators (sparrowhawk Accipiter nisus and domestic cat Felis catus). The birds were startled by moving dummies of these predators and respective control objects, and their risk taking was estimated as their latency to feed after the startle. We found that sparrows responded more strongly (had longer post‐startle feeding latencies) to sparrowhawk attacks than to the control object, and their responses differed between the habitats. First, risk taking of urban birds strongly decreased with age (older birds had longer latencies than young birds), while there was no such age difference in rural birds. Second, young urban birds responded less strongly, while older urban birds responded more strongly to the sparrowhawk than the same age groups of rural birds, respectively. We did not succeed in evoking antipredatory response by simulated cat attacks, because birds responded similarly to the dummy and the control object. Our results support that predation risk, posed at least by avian predators, is different in urban and rural habitats of house sparrows. The increased wariness of older, hence presumably more experienced, urban birds implies that sparrows may be more exposed to predation in cities.     

Predation-Related Costs and Benefits of Conspecific Attraction in Songbirds—An Agent-Based Approach

Songbirds that follow a conspecific attraction strategy in the habitat selection process prefer to settle in habitat patches already occupied by other individuals. This largely affects the patterns of their spatio-temporal distribution and leads to clustered breeding. Although making informed settlement decisions is expected to be beneficial for individuals, such territory clusters may potentially provide additional fitness benefits (e.g., through the dilution effect) or costs (e.g., possibly facilitating nest localization if predators respond functionally to prey distribution). Thus, we hypothesized that the fitness consequences of following a conspecific attraction strategy may largely depend on the composition of the predator community. We developed an agent-based model in which we simulated the settling behavior of birds that use a conspecific attraction strategy and breed in a multi-predator landscape with predators that exhibited different foraging strategies. Moreover, we investigated whether Bayesian updating of prior settlement decisions according to the perceived predation risk may improve the fitness of birds that rely on conspecific cues. Our results provide evidence that the fitness consequences of conspecific attraction are predation-related. We found that in landscapes dominated by predators able to respond functionally to prey distribution, clustered breeding led to fitness costs. However, this cost could be reduced if birds performed Bayesian updating of prior settlement decisions and perceived nesting with too many neighbors as a threat. Our results did not support the hypothesis that in landscapes dominated by incidental predators, clustered breeding as a byproduct of conspecific attraction provides fitness benefits through the dilution effect. We suggest that this may be due to the spatial scale of songbirds’ aggregative behavior. In general, we provide evidence that when considering the fitness consequences of conspecific attraction for songbirds, one should expect a trade-off between the benefits of making informed decisions and the costs of clustering.     

Learning to Choose Among Social Foraging Strategies in Adult House Sparrows (Passer domesticus)

Social foragers may be regarded as being engaged in a producer–scrounger game in which they can search for food independently or join others who have discovered food. Research on the producer–scrounger game has focused mainly on the different factors influencing its evolutionarily stable strategy (ESS) solution, but very little is known about the actual mechanisms that shape players' decisions. Recent work has shown that early experience can affect producer–scrounger foraging tendencies in young house sparrows and that in nutmeg mannikins learning is involved in reaching the ESS. Here, we show that direct manipulation of the success rate experienced by adult sparrows when following others can change their strategy choice on the following day. We presented to live sparrows an experimental regime, where stuffed adult house sparrows in a feeding position were positioned on a foraging grid that included two reward regimes: a positive one, in which the stuffed models were placed near food, and a negative one, in which the models were placed away from food. There was a significant increase in joining behavior after the positive treatment (exhibited by 84% of the birds), but no change after the negative treatment. Further analysis demonstrated that sparrows more frequently used the strategy with which they were more successful (usually joining) and that differences in strategy use were correlated with differences in success. These results suggest that adult birds can monitor their success and learn to choose among social foraging strategies in the producer–scrounger game.     

Causes of reduced clutch size in a tidal marsh endemic

We tested three hypotheses of clutch size variation in two subspecies of the swamp sparrow (Melospiza georgiana georgiana and M. g. nigrescens). Swamp sparrows follow the pattern of other estuarine endemics, where clutch size is smaller among tidal salt marsh populations (M. g. nigrescens) than their closest inland relatives (M. g. georgiana). Our results support predation risk and temperature, but not adult survival, as explanations of this pattern in swamp sparrows. Coastal nests were twice as likely to fail as inland nests, and parental activity around the nest site was positively related to clutch size at both sites. When brood size was controlled for, coastal adults visited nests less often and females vocalized less frequently during visits than inland birds, which may decrease nest detectability to predators. Coastal parents waited longer than inland birds to feed offspring in the presence of a model nest predator, but there was no difference in their response to models of predators of adults, as would be expected if coastal birds possessed increased longevity. Additionally, coastal females laid more eggs than inland females over a single season, following a within-season bet-hedging strategy rather than reducing within-season investment. Coastal territories experienced ambient air temperatures above the physiological zero of egg development more often, and higher temperatures during laying correlated with smaller clutches and increased egg inviability among coastal birds. Similar effects were not seen among inland nests, where laying temperatures were generally below physiological zero. Both subspecies showed an increase in hatching asynchrony and a decrease in apparent incubation length under high temperatures. Coastal individuals, however, showed less hatching asynchrony overall despite higher temperatures. Both air temperatures during laying and predation risk could potentially explain reduced clutch size in not only coastal plain swamp sparrows, but also other tidal marsh endemics.     

Integration of environmental stimuli in the migratory orientation of the savannah sparrow (Passerculus sandwichensis)

Two ‘cue-conflict’ experiments were designed to evaluate the role of (1) solar cues at sunset and stars, and (2) solar cues at sunset and geomagnetic stimuli, in the migratory orientation of the savannah sparrow (Passerculus sandwichensis). A sunset and stars experiment exposed birds in the experimental group to a mirror-reflected sunset followed by an unmanipulated view of stars. Experimental birds shifted their migratory activity in accordance with the setting sun despite exposure to a normal night sky. The sunset and geomagnetism experiment exposed birds in the experimental group to a simultaneous shift in both the position of sunset and the earth's magnetic field. Again experimentals shifted their activity in accordance with the setting sun rather than the artificially shifted magnetic field. Savannah sparrows probaly use stars as celestial landmarks to maintain a preferred direction and do not reorient their activity when exposed to an alternative cue once a direction is established. Moreover, savannah sparrows with experience of migration do not require geomagnetic information in order to use the solar cues available at sunset to select a migratory direction.     

Is Foraging Time Limited During Winter? – A Feeding Experiment with Tree Sparrows Under Different Predation Risk

Small passerines are faced with a trade‐off when foraging during winter. Increasing energy reserves makes them more vulnerable to predators, while a low level of reserves exposes them to a high risk of starvation. Whether small birds under these circumstances are allowed to reduce their foraging activity under increased predation risk, for example in feeding sites more exposed to predators, remains controversial in former behavioural and ecological researches. In this study, we investigated the foraging activity of free‐living Tree Sparrow Passer montanus flocks feeding on an artificial feeding platform. The predation risk perceived by the sparrows was manipulated by placing the platform either close to or far from a bushy shelter. Foraging activity, assessed as cumulative activity of sparrows per unit time on the platform, did not differ between the low‐risk and the high‐risk conditions and did not significantly change during the day. Feeding efficiency, assessed as pecking rate, was not either reduced under the high‐risk condition. Our results suggest that sparrows were forced to feed almost continuously during the day in order to maintain their preferred level of energy reserves. However, several behavioural changes helped sparrows to adopt a safer foraging policy when feeding far from cover, as we found in another study. Altogether, sparrow flocks feeding far from cover decreased the overall foraging time (the time when any sparrow stayed on the platform) by approximately 20% as compared to the near cover condition. A possible way to maintain the same level of foraging activity despite of the reduction in overall foraging time is discussed.     

Group-foraging is not associated with longevity in North American birds

Group-foraging is common in many animal taxa and is thought to offer protection against predators and greater foraging efficiency. Such benefits may have driven evolutionary transitions from solitary to group-foraging. Greater protection against predators and greater access to resources should reduce extrinsic sources of mortality and thus select for higher longevity according to life-history theory. I assessed the association between group-foraging and longevity in a sample of 421 North American birds. Taking into account known correlates of longevity, such as age at first reproduction and body mass, foraging group size was not correlated with maximum longevity, with and without phylogenetic correction. However, longevity increased with body mass in non-passerine birds. The results suggest that the hypothesized changes in predation risk with group size may not correlate with mortality rate in foraging birds.     

Seeing without being seen: a removal experiment with mixed flocks of Willow and Crested Tits Parus montanus and cristatus

This paper tests the hypothesis that foraging site selection reflects a trade-off between the various needs for concealment from predators, to find food, and for the individual to maintain some view of its surroundings. After removal of Crested Tits Parus cristatus (the dominant species in mixed flocks), Willow Tits P. montanus did not decrease their foraging heights as expected but remained in the most exposed parts of young pines. In contrast, after removal of Willow Tits, Crested Tits increased their foraging height from the sheltered lower canopy to sites previously occupied by Willow Tits. When flock size was reduced, the birds maintained the same high levels of vigilance without concealing themselves in dense vegetation. I suggest that flock members may benefit from foraging in sites that afford good anti-predator vigilance.     

Moonlight and the Migratory Orientation of Savannah Sparrows (Passerculus sandwichensis)

Migratory birds might respond to moonlight in at least four ways: (1) a geographical reference for selecting a compass direction, (2) a celestial ‘landmark’ to facilitate maintenance of a preferred heading, (3) a stimulus that distracts migrants and introduces error in compass orientation, or (4) a source of illumination that facilitates nocturnal flight. This study examines the response of migratory savannah sparrows (Passerculus sandwichensis) to moonlight during controlled tests in orientation cages. I found no evidence that savannah sparrows use a lunar compass to select a direction. If savannah sparrows do use the moon as a ‘landmark’ to maintain a direction selected with reference to a different cue, I expected birds to be better oriented on overcast nights when the moon is present than they are when the moon is absent. The results suggest otherwise. Usually, savannah sparrows respond phototactically to the moon by directing their cage activity toward or at a constant angle with respect to the moon's azimuth. Interestingly, the migrant's response to moonlight depended on whether the bird viewed the setting sun earlier that evening.     

Space and habitat use by Red Kites Milvus milvus during winter in the Guadalquivir marshes: a comparison between resident and wintering populations

Differences in habitat selection, diet and behaviour of resident and wintering Red Kites Milvus milvus were studied in Donana National Park, southwest Spain. Adult resident Red Kites roosted at their nests, while immature residents and wintering birds gathered at communal roosts. Individuals remained on average for two consecutive nights (range 1–8) at the same communal roost. Wintering kites spent significantly more time foraging gregariously than residents. The use of the marsh was greater by wintering kites while residents more often used the forest. Wintering birds consumed more goose carrion than the residents. Wintering kites had larger core areas, moved farther from their roost sites to feed and changed foraging areas more frequently than residents. Among residents, adult females had the smallest home ranges (core area, distances travelled and time spent flying). We suggest that Red Kites wintering in Donana occupy the marsh because it provides high food availability, and is vacated by Black Kites M. migrans which exploit this profitable habitat during the breeding season.     

Influence of environmental variables on foraging by juvenile American Robins

ABSTRACT.   Juvenile birds lack the experience of adults and, as a result, are typically less efficient foragers. Environmental factors can influence how birds forage and the outcome of foraging bouts, but few investigators have considered the effects of such factors on the foraging behavior of juveniles. We examined the effects of two environmental factors, sunlight and soil moisture, on the foraging behavior of juvenile and adult American Robins ( Turdus migratorius ). Both factors had a significant effect on robin foraging, with robins more effective at capturing arthropods in the sun and worms in moist soils. However, juveniles were less successful than adults across all conditions. Juveniles were less successful than adults at capturing arthropods and were less efficient at capturing worms. Juveniles captured an average of one worm per minute, whereas adults captured nearly two worms per minute. Additionally, the high failure rates of juveniles (0.44/min) as compared to adults (0.20/min) may be indicative of their inability to choose suitable prey items. Finally, we found that juveniles tended to forage with other robins more than did adults, suggesting that they may use other individuals as cues for locating favorable foraging sites.     

Stomatal and photosynthetic responses to variable sunlight

Most plants experience many fluctuations in sunlight from full sun to shade throughout the day. Under these conditions, stomatal and photosynthetic responses vary dramatically among species depending on water status and growth form. Many herbaceous, fast-growing species rapidly reduce stomatal opening during short-term shade periods. Rapid stomatal closure during shade conserves water, but may also reduce CO₂ uptake. Because periods of alternating sun and shade can reduce accumulative water stress that would otherwise severely curtail carbon gain, some herbs are restricted to habitats with intermittent periods of shade. In contrast to herbaceous growth forms, woody species maintain relatively constant stomatal opening during both sun and shade periods. This results in greater CO₂ uptake, but with greater water loss. These two generalized response patterns for woody and herbaceous species to natural variations in sunlight conflict with conventional ideas of water use and carbon gain based on measurements made under constant light.     

Hazel grouse (Bonasa bonasia) pairs during the nonbreeding season: mutual benefits of a cooperative alliance

During the nonbreeding season (autumn and winter), hazel grouse(Bonasa bonasia) males and females associate as loosely boundpairs, not as strongly bound pairs as previously thought. Thesepairs could be considered cooperative alliances, with each membergaining both direct and indirect benefits. The most importantdirect benefit appeared to be mutual vigilance against predators.This benefit was found at two levels: at the level of arborealfeeding sites, two birds could forage faster and farther fromcover than single birds, and at the territory scale, the pairwas together more in dangerous habitats. By foraging fartherfrom cover, two birds could use about 23% more of the preferredfood trees and 9%–10% more of the food available withina territory than a single bird, partially mitigating the costof having two birds feeding on limited winter food in the samespace. Defending a common territory appeared to be a less importantdirect benefit of the alliance because the members of a pairdid not defend the same territory and often associated withneighboring birds of the opposite sex. A future benefit of thealliance was having breeding partners in the spring; this wasmost beneficial for males, as the sex ratio was male biased.Members of the pair were often apart. At arboreal feeding sites,this separation was perhaps because the most preferred trees,black alders (Alnns glutinosa) with the most staminate catkins,were located in areas with the most cover, where one bird maybe safer from predators than two. Members of pairs in more securehabitats were more often with extrapair birds of the oppositesex. This suggested a trade-off; birds in safe habitats mayhave visited potential breeding partners, but birds in dangeroushabitats may have had to remain together, foregoing this option,to increase their survival probability. This social organizationappeared to be an adaptation to surviving in a heterogeneoushabitat, with some of the winter food located in dense coverand some located in more open and dangerous situations.     

Interspecific variation in extraction of buried seeds within an assemblage of sparrows

Most North American sparrows forage almost exclusively on herbaceous seeds during the winter months. Limited availability of surface seeds forces some birds to scratch for seeds buried beneath soil, snow, or litter. Artificial seed trays were used to test the ability of five different sparrow species to extract seeds buried at different depths in soil. The results suggest three functional groups based on relative scratching ability. Strong scratchers, which included eastern towhee and song and white-throated sparrows, met or exceeded their energetic requirements when foraging on seeds buried at all depths (down to a maximum depth of 1.50–2.25 cm). A weak scratching species, Savannah sparrow, scratched with the same frequency as the strong scratchers, but experienced negative energy budgets when forced to forage on sub-surface seeds. Finally, a non-scratching species, field sparrow, failed to extract any buried seeds. Level of scratching ability may influence foraging efficiency at low resource densities. As a result, interspecific differences in scratching ability may contribute to habitat selection. Strong scratchers may be adapted to foraging near woody vegetation where intense resource competition and abundant litter limit the availability of surface seeds. Weak scratchers, on the other hand, may be forced to feed away from areas where surface seeds are limited. Since woody vegetation serves as a primary source of protective cover in early successional habitats, a tradeoff between foraging efficiency and the risk of predation may promote the local coexistence of species that differ in relative scratching ability and adaptations to evading predators.