Foraging behaviour of nest predators at open-cup nests of woodland passerines

Nest predation patterns and processes cannot be understood without studying the behaviour of predators. I videotaped the behaviour of 22 species of predators at 171 depredated nests of 13 passerine species, in woodland in the Czech Republic. About 32% (60/187) of all events occurred during the night; mammals accounted for 95% (57/60) and 22% (28/127) of nocturnal and diurnal predation, respectively. About 67% (57/85) of mammalian predation, but only 3% (3/102) of avian predation, occurred during night. Multiple predations by the same species were detected in at least 7% (6/82) and 42% (37/88) of nests depredated by mammals and birds, respectively. Martens Martes martes/foina took nest content mostly all at once; birds (mainly Jay Garrulus glandarius) revisited partially depredated nest during 1–4 consecutive days. Martens stayed at the depredated nest about five times longer than Jays. Martens spent similar time at nests with eggs and nestling, while Jays stayed about twice longer at nests with eggs. Mammals consumed eggs always at the nest (23/23), but took nestlings away in at least 48% (31/64) cases. Birds took the eggs and nestling away in at least 31% (18/58) and 76% (71/94) cases, respectively. Predator visits to active nests without taking the content, repeated partial predation and revisitation of previously depredated nests suggest an effect of memory on predator’s foraging behaviour.     

人工巢箱条件下两种山雀鸟类的同域共存机制

种间竞争会导致鸟类对自身的生存策略进行多方面调整,将多种因素结合起来分析不同鸟类的同域共存机制具有十分重要的意义.2009-2011年的3-7月,在辽宁省东部山区以悬挂人工巢箱的方法招引杂色山雀(Parus varius)和大山雀(Parus major),通过比较这两种鸟类在繁殖时间、繁殖参数和巢址选择上的差异,分析其繁殖期的栖息地选择及繁殖对策,探讨这两种鸟类同域共存的机制.结果表明,生态位的部分分离是大山雀和杂色山雀能够长期共存的基础.两者都是在繁殖期到来立即开始繁殖,均出现两次繁殖高峰,采用消减窝卵数的对策来适应环境质量的下降.对繁殖参数的分析表明,两种鸟类的繁殖对策都为k-选择,但大山雀略偏向r端,大山雀倾向高窝卵数、低繁殖成功率的繁殖策略;杂色山雀倾向低窝卵数、高繁殖成功率的繁殖策略.两种鸟类都选择在林龄较长的森林中筑巢,但是大山雀较注重巢址的安全性能,巢距地面较高,周围乔木密集;杂色山雀较为注重植被类型的选择,巢址多选择在植被多样、灌木生长茂盛的针阔混交林中,推测可能与杂色山雀的食物构成有关.     

Seasonal niche-tracking behaviour of two partially migratory passerines

Migratory birds spend periods of the year in different locations as a response to seasonal changes in environmental suitability. They are classified as either ‘niche-trackers’ or ‘niche-switchers’, depending on whether they track or switch environmental conditions throughout the year. However, the relationship between these strategies and their migratory behaviour is still unclear. Here we examine whether migratory European Robins Erithacus rubecula and Eurasian Blackcaps Sylvia atricapilla track environmental conditions between breeding and wintering areas and whether their behaviour differs from that of their sedentary counterparts. We used ringing data for both migratory and sedentary individuals wintering sympatrically in the Iberian Peninsula to assess the environmental conditions relating to their seasonal distributions. We explored seasonal niche-tracking using two multivariate analyses with alternative sets of predictor variables (landscape and climate) to generate different environmental scenarios. Our results show that migratory individuals track similar climatic conditions throughout their seasonal distributions, whereas sedentary birds cope with great variation in climate over the course of the year. In addition, migratory birds show less seasonal overlap in the landscape structure of their chosen habitats compared with sedentary individuals. These results suggest that there is a trade-off between, on the one hand, the cost for migrants of travelling long distances and, on the other, the increased flexibility required by sedentary birds if they are to tolerate a wider suite of environmental conditions within their permanent ranges. Given that sedentary populations in the southern Iberian Peninsula seem to be linked to populations of migratory individuals of these two species that started to move northwards after the last glacial cycle, the observed patterns suggest that migrant birds represent a fraction of the southern population that is specialized in the exploitation of a narrower range of environmental conditions.     

Intraseasonal patterns in shorebird nest survival are related to nest age and defence behaviour

Nest survival may vary throughout the breeding season for many bird species, and the nature of this temporal variation can reveal the links between birds, their predators, and other components of the ecosystem. We used program Mark to model patterns in nest survival within the breeding season for shorebirds nesting on arctic tundra. From 2000 to 2007, we monitored 521 nests of five shorebird species and found strong evidence for variation in nest survival within a nesting season. Daily nest survival was lowest in the mid-season in 5 of 8 years, but the timing and magnitude of the lows varied. We found no evidence that this quadratic time effect was driven by seasonal changes in weather or the abundance of predators. Contrary to our prediction, the risk of predation was not greatest when the number of active shorebird nests was highest. Although nest abundance reached a maximum near the middle of the breeding season, a daily index of shorebird nest activity was not supported as a predictor of nest survival in the models. Predators’ access to other diet items, in addition to shorebird nests, may instead determine the temporal patterns of nest predation. Nest survival also displayed a positive, linear relationship with nest age; however, this effect was most pronounced among species with biparental incubation. Among biparental species, parents defended older nests with greater intensity. We did not detect a similar relationship among uniparental species, and conclude that the stronger relationship between nest age and both nest defence and nest survival for biparental species reflects that their nest defence is more effective.     

Correlation between two components of parental investment: nest defence intensity and nestling provisioning effort of willow tits

Nest defence intensity and nestling provisioning effort of female willow tits (Parus montanus) were significantly correlated at the end of nestling period: well-fed young were defended most intensely. Increased effort was rewarded, since broods with the highest female per-offspring provisioning rates were the most likely to produce local recruits. This suggests that the feeding ability is an important cue for parental investment decisions, at least in a species like the willow tit which has adopted the clutch adjustment strategy. Thus, the most valuable broods would not necessarily be the largest ones, but the ones in which the original number of young could be fed most adequately. However, no associations were found between the level of parental effort and offspring weight, size or condition, nor did the broods producing recruits differ from other broods in timing of breeding or number and size of offspring. The female behaviour may suggest that they invest the most time, energy and risk in the young whose chances of joining the winter flock are the best. The first well-fed young also gain an advantage of prior residency in joining the flock. The first to join normally obtain higher social status, and hence better winter survival, than latecomers. The corresponding patterns in male parental investment behaviour were weak or absent, which suggested that the male effort was affected by the female behaviour. Males seemed to invest in nestling provisioning in such a way as to supplement the female effort. During nest defence action males also seemed to invest in protection of females against predation.     

Nest intruders, nest defence and foraging behaviour in the Black-and-white Casqued Hornbill Bycanistes subcylindricus

The female Black-and-white Casqued Hornbill Bycanistes subcylindricus uses mud to seal herself into a nest cavity and remains there until the nesting attempt has ended. The male hornbill is solely responsible for food provisioning and external defence of the nest. Data on hornbill nests in a Ugandan rain forest were used to test Martindale's (1982) model of nest defence and central place foraging. As predicted, nest guarding by the resident male hornbill during intrusions by conspecifics altered foraging patterns; the resident male foraged closer to the nest, made a larger number of shorter visits, and brought smaller food loads. There was a significant change in size composition of fruits brought before, during, and after intrusions. For short intrusions, the volumes of food brought per unit time did not change. However, if intrusions lasted for days or weeks, food delivery rates declined. Nest-sealing by the Black-and-white Casqued Hornbill appears to function primarily to protect the nest from conspecifics rather than from interspecific predators.     

Nest defence,enemy recognition and nest inspection behaviour of experimentally parasitized Reed Warblers Acrocephalus scirpaceus

Capsule Reed Warblers in a regularly parasitized population do not recognize Cuckoo Cuculus canorus as a special enemy and do not change their behaviour at nest immediately after being parasitized.

Aims To assess if an intruder near the nest influences the behaviour of the Cuckoo host.

Methods Host responses to Cuckoo, control Pigeon dummies and human intruder were observed. Host behaviour at 71 nests was video-recorded for 30 minutes at four experimental groups of nests: Cuckoo dummy, Cuckoo dummy + Cuckoo egg, Pigeon dummy, human intruder.

Results Reed Warblers did not respond differently to the Cuckoo and the control species. The experimental procedure had no significant effect on the behaviour of hosts during the study period. We were unable to find any differences in the time spent at the nest, clutch inspection behaviour and nest defence behaviour between morning and afternoon experimental groups. Our results do not support the hypothesis that afternoon laying by the Cuckoo is maintained by a selection pressure from the host. We observed no ejection or egg-pecking during the 30-min period after the experimental parasitism.

Conclusions Low aggression and non-specificity of host responses in our study area are in line with the fact that the Reed Warbler is an intermediate rejecter of Cuckoo eggs as expected from the spatial habitat structure hypothesis.     

Supplemental food alters nest defence and incubation behaviour of an open‐nesting wetland songbird

Climate‐driven increases in spring temperatures are expected to result in higher prey availability earlier in the breeding season for insectivorous birds breeding in wetland habitats. Predation during the incubation phase is a major cause of nesting failure in open‐nesting altricial birds such as the Eurasian reed warbler. The nest predation rate in this species has recently been shown to be substantially reduced under conditions of experimentally elevated invertebrate prey availability. Food availability near the nest may be an important determinant of adult incubation and nest defence behaviours during the incubation period. We used two experimental studies to compare incubation behaviour and nest defence in food‐supplemented and unsupplemented adult Eurasian reed warblers during the incubation phase. In the first study we measured nest defence behavioural responses to a taxidermic mount of a native predator (stoat Mustela erminea). In the second study we used temperature loggers installed in nests to measure breaks in incubation as a measure of nest vulnerability. Food‐supplemented birds responded aggressively to the presence of a predator more quickly than those in the unsupplemented group, suggesting they are closer to their nest and can more quickly detect a predator in the vicinity. Food‐supplemented birds also had shorter breaks in incubation (both in terms of maximum and mean off‐bout durations), presumably because they were foraging for shorter periods or over shorter distances from the nest. This study therefore identifies the behavioural mechanisms by which changes in food availability may lead to changes in nest survival and thus breeding productivity, in open‐nesting insectivorous birds.     

Linking anti-predator behaviour and habitat quality: group effect in nest defence of a passerine bird

In habitats where the density of breeding individuals is higher, breeding success has been shown to increase with the number of close conspecific and heterospecific neighbours. However, the mechanisms linking habitat quality, group size of prey individuals and offspring defence are poorly known. In this field study, we examined the relationships between habitat quality and parental nest defence behaviour in the pied flycatcher (Ficedula hypoleuca). We found that mobbing is more intense in unmanaged forests where birds breed in more dense and diverse communities than in heavily managed young forests where heterospecific densities are lower. We also found that the mobbing activities of pied flycatchers breeding in unmanaged mature boreal forests attracted more neighbouring prey individuals than in nearby managed forests. This study shows that habitat quality-mediated effects might be responsible for the decreased group size of mobbing birds in managed forests, which may lead to less effective communal defence.     

Regional variation in seasonality affects migratory behaviour and life-history traits of two Mediterranean passerines

Migratory birds may improve fecundity by moving to seasonal breeding areas, but may also suffer higher mortality rates as a cost of movement. However, the covariation among seasonality, migratoriness and life histories should change between species if their ecological features affect site-tenacity, survival or fecundity. In frugivorous birds, for instance, wandering in search of fruits may trigger broader migrations than territorial defence, and also may improve nonbreeding survival by preventing food shortages that eventually happen in discrete territories. We studied the variation in spatio-temporal distribution, life expectancy and fecundity in robins (Erithacus rubecula) and blackcaps (Sylvia atricapilla) distributed in three Iberian regions with a low, mid or high degree of environmental seasonality. In the Iberian Peninsula, robins and blackcaps are the two most intensive frugivores in winter; however, robins are territorial while blackcaps track fruit abundance among habitat patches. In the most seasonal area, robins and blackcaps decreased abundance in winter and showed a lower breeding-site tenacity, a shorter life expectancy and a larger clutch size. However, blackcaps tended to be more migratory than robins in all regions. Despite their stronger migratory behaviour, blackcaps showed a longer life expectancy and a smaller clutch size than robins in all regions. In robins, winter territoriality could decrease nonbreeding survival but also improve fecundity, because survivors are dominant and hence more efficient breeders. In blackcaps, however, the use of the most profitable habitat patches could improve nonbreeding survival, thereby allowing a similar recruitment than in robins with a lower reproductive investment. These results support that regional-scale changes in seasonality may affect migratory behaviour and hence the tradeoff between reproduction and survival in birds, doing so differently depending on the idiosyncrasy of each species.     

Temporal differences in food abundance promote coexistence between two congeneric passerines

Many related species share the same environment and utilize similar resources. This is surprising because based on the principle of competitive exclusion one would predict that the superior competitor would drive the other species to extinction; coexistence is only predicted if interspecific competition is weaker than intraspecific competition. Interspecific competition is frequently reduced by differential resource use, resulting in habitat segregation. In this paper, we use the closely related collared and pied flycatcher to assess the potential of habitat differences to affect interspecific competition through a different mechanism, namely by generating temporal differences in availability of similar food resources between the two species. We found that the tree species composition of the breeding territories of the two species differed, mainly by a higher abundance of coniferous species around nest-boxes occupied by pied flycatchers. The temporal availability of caterpillars was measured using frass traps under four deciduous and two coniferous tree species. Deciduous tree species showed an early and narrow peak in abundance, which contrasted with the steady increase in caterpillar abundance in the coniferous tree species through the season. We subsequently calculated the predicted total caterpillar biomass available in each flycatcher territory. This differed between the species, with biomass decreasing more slowly in pied flycatcher territories. Caterpillar biomass is strongly correlated with the reproductive success of collared flycatchers, but much less so with pied flycatchers. However, caterpillar availability can only partly explain the differences in seasonal decline of reproductive success between the two species; we discuss additional factors that may contribute to this species difference. Overall, our results are consistent with the suggestion that minor habitat differences between these two species may contribute to promoting their coexistence.     

Species-specific song convergence in a moving hybrid zone between two passerines

Moving hybrid zones are receiving increasing attention. However, so far little is known about the proximate mechanisms underlying these movements. Signalling behaviour, by individuals engaged in interspecific sexual and aggressive interactions, may play a crucial role. In this study, we investigated song variation within a moving hybrid zone between two warblers, Hippolais polyglotta and H. icterina . In these species, song is involved in interspecific territoriality and, probably, in mixed pairings. We showed that allopatric populations of the two species are clearly acoustically differentiated. However, interspecific differences faded out in sympatry as a result of an overall pattern of convergence. Unexpectedly, the two species converged for different song parameters, namely temporal parameters for H. icterina and syntax for H. polyglotta . Hybridization and interspecific competition could explain convergence in H. icterina . Instead, in H. polyglotta we suggest that local adaptation to habitat and interspecific learning might contribute to convergence. We particularly stress that cross-species learning, by maintaining high levels of interspecific interactions, may influence the movement of the zone.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 507–517.     

Long-term familiarity promotes joining in neighbour nest defence

Familiarity plays an important role in the evolution of sociality and cooperation. Familiar individuals may gain a reputation for participating in, or defecting from, cooperative tasks. Previous research suggests that long-term familiarity with territorial neighbours benefits breeders. We tested the hypothesis that great tits (Parus major) are more likely to join in neighbours' nest defence if those neighbours are familiar from the previous year. We show that neighbours that shared a territory boundary the previous year are more likely to join their neighbours' nest defence than neighbours that did not share a boundary before. Closer neighbours did not differ from distant neighbours in their latency to join. For familiar neighbours that joined, there was no difference in call rate in relation to whether one or both members of the focal pair were familiar. First-time breeders (by definition unfamiliar) did not join each other's nest defence. This is the first evidence of a relationship between familiarity and joining in nest defence. Such direct benefits of familiarity may have important implications in the evolution of sociality.     

Repeatability of nest predation in passerines depends on predator species and time scale

It has been proposed that some specific locations of bird's nests have higher intrinsic chances of being depredated than other locations. This predicts that fates of consecutive nesting attempts at the same site should be repeatable. We used 20 pairs of old thrush nests to simulate repeated nesting attempts at the same sites, both within and between breeding seasons (n=40  sites×2  trials×2  years=160). Each nest was monitored by a camera to record multiple predation events and to identify predators. Predation by all predator species was repeatable during a 15-day trial. Predation by principal predators (jay Garrulus glandarius , marten Martes martes / foina ) and total predation (all species combined) was not correlated within pairs of simultaneously exposed nests or within samples of nests from particular study plot, and not repeatable for individual nests between-trials or between-years. These findings suggest short-term effect of predator memory causing revisitation of previously depredated nests during a current nesting trial (all predators); do not support an effect of nest site features on multiple nest discoveries and/or an effect of nest location on repeated random encounters with the same nest (principal predators). Long-term repeatability and correlation within pairs of simultaneously exposed nests was detectable only in occasional predators (great spotted woodpecker Dendrocopos major , possibly also squirrel Sciurus vulgaris ), which suggests effect of nest location combined with site fidelity and individual foraging specialization of these predators. We conclude that repeatability of nest predation depends on the time scale considered and the local predator community. We caution against spurious findings of repeatable nest predation resulting simply from statistical properties of correlation in binary data (nest fates).     

Parental behaviour in Copris lunaris (Coleoptera, Scarabaeidae): care and defence of brood balls and nest

Abstract. 1. Nesting female beetles righted brood balls (so as to replace the egg or larva in the uppermost position) and repaired damaged balls. This behaviour required the presence of an egg or larva in the ball, or of a short-lasting material found just after oviposition. The shape of the ball was also a righting stimulus since artificial ellipsoids were stood on end.
2. Balls containing dichloromethane extracts of C.lunaris brood were righted and repaired. Eggs and larvae of several other Scarabaeidae did not release these responses but were destroyed.
3. Righting behaviour was released when brood was absent from the top of the ball. The beetle then crawled vertically downwards and, if it encountered the displaced apex, a novel rolling action followed which automatically turned the ball towards the correct position.
4. An opening made in the nest was repaired with soil excavated from the chamber floor. Clunaris adults and Aphodius fossor larvae were attacked if they were encountered in the nest.     

Effectiveness of nest defence in the Acadian Flycatcher Empidonax virescens

We used presentations of models to determine the effectiveness of nest defence in the Acadian Flycatcher Empidonax virescens against a nest predator (Blue Jay Cyanocitta cristata ) and a brood parasite (Brown-headed Cowbird Molothrus ater ). Principal components analysis (PCA) of four component variables of nest defence (call rate, swoop rate, closest approach and number of adults) generated a measure of overall nest defence (aggression). We determined effectiveness of defence by looking for correlations between measures of defence and measures of nest success (nest predation and brood parasitism). We also determined whether nest defence increased with clutch size, nestling age and time in the breeding season. Defence against model Brown-headed Cowbirds did not correlate with levels of parasitism, clutch size, age of young or time of breeding. There was, however, a strong, but insignificant, trend for nests with high levels of all measures of defence to suffer less from brood parasitism. Aggression, vocalization rate, closest approach and number of adults defending against models of predatory Blue Jays correlated positively with nesting success during the egg stage but not the nestling stage of the nesting cycle. Aggression, vocalization rate, closest approach correlated with clutch size and age of the brood. These results suggest that nest defence can effectively deter nest predators, but may be less effective against brood parasites. Different behavioural components of nest defence may work at different stages of the nest cycle and against different nest predators. The components of nest defence that correlated with nest success also correlated with clutch value, a result consistent with hypotheses on the evolution of nest defence.