Parental Investment Theory and Nest Defence by Imperial Shags: Effects of Offspring Number,Offspring Age,Laying Date and Parent Sex

Nest defence is a common form of parental care employed by birds to improve the survival of their offspring. Theory predicts that parents should adjust their nest defence according to the value of the brood at stake, defending more intensively broods with high survival and reproductive prospects. We evaluated the influence of offspring number, offspring age, laying date and parent sex on nest‐defence intensity (NDI) of the Imperial Shag Phalacrocorax atriceps, a sexually dimorphic seabird with seasonal decline in offspring survival and very limited renesting potential. We also evaluated whether NDI was correlated within pairs and whether NDI of both members of the pair was correlated with incubation and breeding success. To elicit defensive behaviour, we simulated predation attempts using a Kelp Gull Larus dominicanus model. As predicted by theory, NDI was positively correlated with the number of offspring in the nest and offspring age. NDI during chick rearing was higher than that at early and late incubation, while no differences were found between incubation stages. Contrary to our prediction, we did not find differences in NDI according to laying date. NDI for males was higher than females, while NDI was also positively correlated within pairs. NDI was not statistically related to incubation or breeding success. These results suggest that other factors, such as laying date or parental quality and age, play a much larger role in determining the outcome and productivity of a nesting attempt. Our results provide partial support for parental investment theory; while parental defence increased with brood value according to offspring number and age, parental defence was not related to laying date, a factor strongly affecting offspring survival and recruitment probabilities in this species.     

Nest defence in the meadow pipit <Emphasis Type="Italic">Anthus pratensis</Emphasis>: the influence of renesting potential

The reproductive value hypothesis predicts that the level of nest defence is determined by the expected chance of offspring to survive until reproduction, and by the reproductive potential of the parents. Rates of survival from one breeding season to the next are low in small passerines, and their residual reproductive potential strongly declines as the current breeding season terminates. Therefore, we can expect that parents which have only one breeding attempt per season should defend their nests more intensively than parents with a possibility to renest. We studied nest defence in populations of meadow pipit (Anthus pratensis) breeding in Norway and the Czech Republic, differing in renesting potential. To simulate the threat from a predator, we placed a stuffed stoat (Mustela erminea) first 5 m and then 1 m away from a nest with nestlings. Parents increased or kept nest defence constant when the stoat approached their nests in Norway and, during a breeding season shortened by severe weather, in the Czech Republic (when renesting potential was limited). Parents decreased nest defence when the stoat approached the nest during “normal” breeding seasons in the Czech Republic (when renesting was common). These findings give support to the reproductive value hypothesis.     

Nest Defence of Nesting Chinstrap Penguins (Pygoscelis antarctica) against Intruders

We tested some predictions of parental investment theory by studying the aggressive behaviour of colonial nesting chinstrap penguins (Pygoscelis antarctica) against human intruders into their nesting territories. We tested for differences in the aggressive behaviour of penguins according to offspring age (eggs vs. chicks), offspring number, nest location in the colonies (central vs. peripheral) and sex. Offspring age was the main factor influencing nest defence, although nest location and sex were also important. Chicks were defended more strongly than eggs, in accordance with changes in the reproductive value of offspring, and this increase in aggressiveness was not related to revisitation of the same individuals. The level of aggression of penguins breeding in central sites was higher than that of peripheral birds, a difference that could be due to the lower residual reproductive value of central-nesting, probably older, birds. The stronger aggressiveness of males could be due to a combination of factors related to sexual selection and life-history traits. Offspring number did not affect the level of nest defence.     

Reproductive tactics of the Asian cichlids of the genusEtroplus in Sri Lanka

Synopsis The Asian cichlids,Etroplus maculatus (the orange chromide) and E. suratensis (the green chromide) in Sri Lanka reproduce twice during the year when water conditions are favorable for nest construction and maintaining visual contact with offspring. These are the drier premonsoonal and monsoonal seasons when water turbidity decreased and salinity increased. When breeding in isolation orange chromide pairs selected dense vegetation where nests were camouflaged. During the peak breeding cycle (July) orange chromide pairs selected sparse vegetation for nesting as a compromise between survival of young and availability of adult food. These areas were also occupied by foraging non-breeding conspecifics which increased the threat of cannibalism of offspring. Under these pressures most orange chromides nested in colonies which helped decrease both actual and attempted cannibalism. The sympatric green chromide does not forage during nesting and nest site selection was determined mainly by factors favoring offspring survival. Biparental care is exhibited by both species. One member of an orange chromide pair stands guard over offspring while the other leaves the territory to forage — their roles are reversed every few minutes. The total parental investment is equivalent to the full investment of a single parent. Monogamy appears to be maintained by their metabolic constraints. In the green chromide both parents are vigilant over offspring and neither forages thus spending twice as much time in parental investment.     

Temporal Variation in Decisions about Parental Care in Bluegill, Lepomis macrochirus

Parental investment theory states that an individual will trade‐off present and future reproductive potential to maximize lifetime reproductive success. Only when parental care is costly in terms of reduced future reproductive potential should individuals be sensitive to changes in the value of current offspring and adjust their care. Here, we examine temporal variation in parental care decision‐making in bluegill (Lepomis macrochirus), in which care is provided by males called ‘parentals’. Previous research has shown that parentals that nest early in the breeding season are in higher energetic condition than those that nest later, and early nesting males appear not to pay an opportunity cost to their care in terms of reduced future reproductive potential. Early nesting males also may have higher paternity in their broods than later nesting males. To examine the parental care decisions made by early and mid‐season nesting parentals, we experimentally reduced males’ perceived paternity by swapping eggs between nests. We found that experimental males that nested early in the breeding season adjusted their brood defence behaviour similarly to control males, which had sham egg swaps performed. Conversely, experimental males that nested mid‐season significantly decreased their brood defence behaviour after the manipulation as compared with control males. Thus, unlike mid‐season nesting males, early nesting males appear relatively insensitive to changes in brood value (paternity), possibly because early nesting males pay little cost in terms of reduced future reproductive potential to providing full care or because these males have a predisposition to high paternity.     

Flexible responses to stage‐specific offspring threats

When caring for their young, parents must compensate for threats to offspring survival in a manner that maximizes their lifetime reproductive success. In birds, parents respond to offspring threats by altering reproductive strategies throughout the breeding attempt. Because altered reproductive strategies are costly, when threats to offspring are limited, parents should exhibit a limited response. However, it is unclear if response to offspring threat is the result of an integrated set of correlated changes throughout the breeding attempt or if responses are a flexible set of dissociable changes that are stage‐specific. We test these hypotheses in a system where house wrens (Troglodytes aedon) compete for nesting cavities with Carolina chickadees (Poecile carolinensis) by usurping and destroying their nests during the early stage of the breeding attempt (the egg stage). Due to the specificity of the house wren threat, we can test whether parental responses to an offspring threat show flexibility and stage specificity or if parental strategies are an integrated and persistent response. We monitored nests in a natural population to compare life history traits of chickadees nesting in boxes that were in the presence of house wrens to chickadees nesting in boxes that did not overlap with house wrens. Carolina chickadees that nested near house wrens laid significantly smaller clutch sizes (early change in reproductive strategy) but did not alter nestling provisioning or nestling stage length (late change in reproductive strategy), suggesting that chickadees respond in a flexible and stage‐specific manner to the threat of house wrens. By responding only when a threat is highest, parents minimize the cost of antithreat responses. Our study suggests that parents can respond in subtle and nuanced ways to offspring threats in the environment and specifically alter reproductive behaviors at the appropriate stage.     

Cooperative nest‐defence behaviour and territory quality in a resident and socially monogamous passerine

Nest defence is a fundamental aspect of parental care in secondary cavity‐nesting birds, and predation or competition for nesting sites can involve different defensive behaviours. Because habitat quality determines breeding success, we were interested in whether breeding pairs of the Eurasian nuthatch, Sitta europaea, established in more favourable environment also manifest higher probability of cooperative behaviour during their nest‐site defence. To explore this relationship, we quantified behavioural displays of both parents and analysed activity budget ethogram data from simulated territorial intrusions performed in the chick‐feeding phase with one conspecific and two different heterospecific stimuli (dummies of nuthatch, starling and woodpecker). We found that paired individuals shared their roles during nest‐site defence to a considerable extent. Males had a significantly higher number of attacks on intruders than females, and females performed more threat displays and controls of the brood than males. Multinomial analysis of the cooperative behaviour suggested that pairs in a high‐quality territory had higher probability of reciprocal substitution of different roles towards a balance between attacks, threat displays and nest controls. Contrary to this, pairs in a low‐quality territory had less likely pairwise combinations of simultaneous behavioural states that are associated with effective nest‐site defence. The difference in response probability according to territory quality was, however, highly variable in view of the stimulus that was used in simulated territorial intrusion. Because individual roles and the complex behavioural repertoire of pairs altered in response to territory quality and potential nest‐site competitor or brood predator, our results suggest that the cooperative nest‐defence behaviour could be linked to the breeding success of this year‐round territorial species living in a heterogeneous forest habitat.     

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.     

Interactions between common buzzard Buteo buteo and goshawk Accipiter gentilis: trade-offs revealed by a field experiment

I examined the behavioural interactions between common buzzard Buteo buteo and goshawk Accipiter gentilis and their effects on buzzard breeding success and brood defence with a two-year field experiment using dummies and playback calls. A priori I showed through an extensive nest site analysis that there is considerable nesting habitat overlap between the two species and hence potential for interspecific competition for prime nesting habitat. Buzzards had a significantly lower breeding success when presented with a goshawk dummy compared to control broods but there was no effect of buzzard dummies on reproductive success. Buzzards failing with their breeding attempt tended to select another nest site while successful buzzards more frequently used the same nest again. Buzzard pairs were less often attacked by common crows Corvus corone while exposed to goshawk dummies compared to buzzard dummies. The decision to desert a nest seems to be a trade-off between predation risk on the one hand and protection against crows on the other. Goshawks proved far more aggressive against an intraspecific dummy than buzzards. Buzzards adjusted their level of brood defence against both intra- and interspecific dummies according to the age of offspring but not offspring number, with an increasing brood defence level with increasing offspring age. Thus the behaviour of buzzards towards goshawks is a result of a complex system of trade-offs between predation risk, competition for prime nesting habitat and protection from crows on which brood value acts as a temporal modifier.     

Responses of naive yellow warblers to a novel nest predator

Naive parental yellow warblers, Dendroica petechia, were tested with a mounted eastern grey squirrel, Sciurus carolinensis, a novel predator on the study site, during the nest-building, egg-laying, incubation and nestling stages of the nesting cycle. Testing parents only once avoided the possibility that the nest defence behaviour of the parents was modified by reinforcement and loss of fear. In 96% of 80 trials, the female parent responded first to the model. Males rarely showed nest defence behaviour. Female nest defence as measured by defensive vocalizations (i.e., ‘chipping’), closest and modal distance of approach to the model, distraction displays, strikes, and close passes or hovers, did not increase significantly over the breeding season. ‘Chipping’ frequency by both sexes tended to increase after the incubation period but not significantly so. Distraction displays increased significantly only between nest-building and nestling stages. Age, clutch or brood size, and nest success were not correlated with the intensity of nest defence. Females defending exposed nests gave more distraction displays than those defending cryptic nests.     

海拔梯度变化对黄缘蜾蠃筑巢结构及相关生物学特性的影响

黄缘蜾蠃(Anterhynchium flavomarginatum)是一种以鳞翅目幼虫为食的独栖性捕食昆虫,在生态系统中提供了重要的生物防治功能。2018—2019年,在广东车八岭国家级自然保护区通过人工巢穴收集了不同海拔梯度下(332—1007 m)该蜂的筑巢巢管,系统研究了黄缘蜾蠃的筑巢特征(巢管内径、巢室长度、巢室数量)以及该蜂的子代雌雄数和体型大小随海拔梯度的变化趋势。结果表明：海拔主要影响的是黄缘蜾蠃对巢管内径大小的选择,以及对前庭长度的分配,同时,黄缘蜾蠃对巢管内径的选择可以影响到后代性别比,这种策略主要体现在亲本对雌性子代的投资,而不是雄性子代。研究结果表明黄缘蜾蠃在不同海拔梯度下可根据外界环境压力调整其营巢和生殖策略,以提高种群对环境的适应性。     

When do altricial birds reach maximum of their brood defence intensity?

It has been suggested that the brood defence by parents of altricial birds should increase during the breeding attempt until the young depart from the nest. The two proximate hypotheses provide alternative predictions about the peak of brood defence intensity: (1) the vulnerability hypothesis predicts a rapid rise in brood defence after hatching of the chicks, with maximum defence intensity just before fledging and strong decline afterwards; (2) the feedback hypothesis predicts that brood defence intensity will, after a rapid rise, reach a plateau at the end of the nestling period and early after fledging and then slowly decline. I compared brood defence behaviour of altricial meadow pipit (Anthus pratensis) breeding in the Czech Republic during the late nestling stage and during the fledging time. A stuffed stoat (Mustela erminea) was placed 5 m from a meadow pipit nest and the defence behaviour of parents was recorded for 10 min from a hide. Brood defence intensity was higher during the fledgling time than during the late nestling stage, and this trend was more evident in males than in females. Regardless of the proportion of already fledged chicks and those still present in the nest, brood defence did not significantly decrease during the fledgling time in males or females. The results do not agree with the predictions of the vulnerability hypothesis and support the predictions of the feedback hypothesis.     

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.     

Does variation in female body size affect nesting success in Dawson’s burrowing bee,Amegilla dawsoni (Apidae: Anthophorini)?

Abstract.  1. Females of Dawson's burrowing bees vary in body weight over a twofold range. Despite the potential for differences in body weight to affect several aspects of the competition among nesting females, no clear advantages were documented for larger females.
2. Nesting females were not consistently larger than emerging females, nor was there a consistent relationship between body size and the weight of pollen and nectar carried to the nest on provisioning trips. At one nesting location, larger females did not produce larger pre-pupal offspring nor did they produce offspring at a faster rate than their smaller nesting companions.
3. In addition, large body size was not associated with greater success in nest defence despite the fact that nesting females regularly encountered intruders in their burrows. Residency, not body size, determined the outcome of almost all contests for control of a nest burrow. The absence of a large body size effect here appears to stem from an intruder strategy designed to enable nest-searching females to acquire burrows that had been abandoned through death or dispersal of their original owners rather than securing existing nests through an aggressive takeover strategy.
4. Thus, although large body size conveys significant fitness advantages to males, this attribute does not promote female success in either provisioning or defending their nests.     

Enemy Recognition of Reed Warblers (Acrocephalus scirpaceus): Threats and Reproductive Value Act Independently in Nest Defence Modulation

Organisms should respond more aggressively towards species perceived as a danger to their offspring, but intensity of defence may be gauged by the value of current offspring weighed against the value of future reproductive opportunities. We tested whether defensive responses of nesting reed warblers (Acrocephalus scirpaceus) are the result of an interaction effect between the type of stimulus confronted and the value of the warbler’s nesting attempt. We quantified the ability of reed warblers to discriminate among brood parasites, nestling predators and non‐threatening species at different stages of the breeding cycle. We also determined whether variables that influence the value of offspring, such as time of season, size and age of clutch or brood, and time of day and number of visits to the nest, explain variation in the intensity of defence recorded during the egg and nestling stages. Responses to the three stimuli differed significantly, as reed warblers consistently directed their mobbing calls and attacks towards parasites, whereas they were less conspicuous when confronted with models of predators. Reed warblers modulated their responses towards each stimulus in accordance with the threat each posed at a specific nesting stage, whereas they were not affected by other variables relative to their reproductive potential. The churr call, however, was uttered independently of the stimulus, as it was triggered by the mere presence of nestlings in the nest.     

Breeding dispersal of Northern Flickers Colaptes auratus in relation to natural nest predation and experimentally increased perception of predation risk

After nest predation, breeding dispersal can be an effective strategy to avoid local nest predators. Furthermore, encounters with predators at a nest during the pre-laying stage may be used by parents to judge future risk, such that they may abandon a nest when a nest predator has been encountered. We studied whether the between- and within-year breeding dispersal of Northern Flickers Colaptes auratus was dependent upon the outcome of the previous nesting attempt. We also tested whether pairs presented with a model predator prior to egg-laying were more likely to abandon their nests than were pairs presented with a control model. Between years, males moved significantly further after having their nest depredated than did successful males, and females showed the same trend. However, these movements did not result in greater reproductive success. More pairs switched sites within years after having their nest depredated, but those that remained and those that moved had equal subsequent nest success. Stressful encounters with predators involving nest defence may trigger dispersal both between and within years, although reproductive benefits are unclear. The proportion of pairs abandoning nests did not differ between parents presented with control or predator models, suggesting that a single encounter with a predator is not a sufficient deterrent against continued use of a particular nest.     

Habitat quality as a predictor of spatial variation in blue tit reproductive performance: a multi-plot analysis in a heterogeneous landscape

Vertebrate studies have rarely investigated the influence of spatial variation in habitat richness on both short-term (breeding) and long-term (offspring recruitment) reproductive performance using simultaneously multi-patch, multi-habitat type and multi-year approaches at landscape level. Here we present results of such an approach using the influence of two oak tree (Quercus ilex, Q. humilis) species on reproductive performance in Corsican blue tits (Parus caeruleus ogliastrae) as a model system. We found that blue tits breeding in rich broad-leaved deciduous patches consistently laid eggs earlier in the season, and produced larger clutches and more fledglings of higher quality, than those breeding in poor evergreen patches. Also, parents, especially males, were in better physical condition in the broad-leaved deciduous than in the evergreen patches. Surprisingly, estimates of long-term effects of reproduction, such as recruitment rates of locally born offspring, did not differ between the two habitat types. Our results suggest that short-term breeding performance and phenotypic quality of both chicks and parents do not necessarily provide reliable information about contributions to following generations at a scale larger than that of the local study plot. Differences in reproductive performance between the two oak habitat types could not be attributed to density-dependent effects, differences in levels of nest predation, or differences in age structure of the birds. We suggest that habitats that are optimal for breeding are not necessarily optimal for survival after the breeding season.     

Costs and benefits of female-biased natal philopatry in the common goldeneye

Sex-biased natal dispersal in long-lived species may resultin interactions between parents and mature young of the philopatricsex. To investigate the evolutionary basis of natal philopatryin a noncooperative species, the common goldeneye Bucephalaclangula, we studied possible costs and benefits of simultaneousbreeding of females and philopatric daughters. We did not find any fitness consequences of a daughter's breeding on their mother'sbreeding in terms of nest-site selection, body weight, clutchsize, hatching date, or hatching success. Our results, therefore,did not support the assumption of the local resource competitionhypothesis, that the natally philopatric sex should be morecostly to a breeding parent. As possible benefits for daughters returning to their natal area, we tested inheritance of nestsites from mothers and explored whether daughters utilize thepresence of their mother by parasitically sneaking into hermother's nest. Daughters' nest-site selection was not associatedwith the presence of their mothers. A comparison between daughtersand control females revealed that daughters chose their nestsite closer to their natal nest than expected by nest-siteavailability alone. Daughters could not expect to inherit anest site from their mother, and we did not find other indicationsof cooperation between relatives either. The mother's clutchsize did not increase in the year breeding with the daughter, indicating daughters do not parasitize their mother's nest.We suggest that benefits such as decreased nest predation riskassociated with nesting close to the natal nest site may beimportant in the natal philopatric behavior of the species.     

Nesting Success in Crimson Finches: Chance or Choice?

In avian systems, nest predation is one of the most significant influences on reproductive success. Selection for mechanisms and behaviours to minimise predation rates should be favoured. To avoid predation, breeding birds can often deter predators through active nest defence or by modifying behaviours around the nest (e.g. reducing feeding rates and vocalisations). Birds might also benefit from concealing nests or placing them in inaccessible locations. The relative importance of these strategies (behaviour vs. site selection) can be difficult to disentangle and may differ according to life history. Tropical birds are thought to experience higher rates of predation than temperate birds and invest less energy in nest defence. We monitored a population of crimson finches (Neochmia phaeton), in the Australian tropics, over two breeding seasons. We found no relationship between adult nest defence behaviour (towards a model reptile predator) and the likelihood of nest success. However, nest success was strongly related to the visibility of the nest and the structure of the vegetation. We found no evidence that adult nest building decisions were influenced by predation risk; individuals that re‐nested after a predation event did not build their nest in a more concealed location. Therefore, predator avoidance, and hence nest success, appears to be largely due to chance rather than due to the behaviour of the birds or their choice of nesting sites. To escape high predation pressures, multiple nesting attempts both within and between seasons may be necessary to increase reproductive success. Alternatively, birds may be limited in their nest‐site options; that is, high‐quality individuals dominate quality nest sites.     

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.