Parent birds assess nest predation risk: influence of cavity condition and avian nest predator activity

Skutch hypothesized that nest predators visually assess parental activities to locate a prey nest, whereas parents modify fitness‐related traits to reduce the probability of nest predation. We examined how cavity condition and parental activity interact with avian nest predators to shape the nest success of two coexisting parid species, marsh tits Poecile palustris and oriental tits Parus minor, breeding in nest‐boxes during the incubation period. Nest‐boxes were manipulated to create a prolonged risk of nest predation (entrance diameter 2.6 cm control vs 5.5 cm treatment) soon after clutch completion. To measure changes in parental behavior, we also simultaneously simulated a pulsed risk of nest predation, using sound playbacks of a coexisting control bird and an avian nest predator. We found that the parent tits merely responded the pulsed risk, presumably due to an environment with high avian nest predator encounters, compared to the prolonged risk. Instead, both species spent more time on vigilance at the nest, only under prolonged risk conditions. The activity of corvids near the nest‐box was higher in the marsh tit than that in oriental tits. This activity was also higher in the treatment nest box than that in the control nest‐box. Nest predation during the incubation period was higher in marsh tits than in oriental tits, presumably due to higher and more plastic vigilance in oriental tits, compared to marsh tits. Our results highlight that the differences in cavity condition and parental activities at the nests of two coexisting non‐excavators may contribute to differential nest predation by attracting avian nest predators.     

An Experimental Study of Predator Recognition in Great Tit Fledglings

Studies of naturally predator-naïve adult birds (finches on predator-free islands) and birds experimentally hand reared in isolation from predators indicate that birds can recognise predators innately; that is, birds show anti-predator behaviour without former experience of predators. To reduce predation risk efficiently during the vulnerable fledgling period, we would predict an innate response to be fully developed when the chicks leave the nest. However, 30-day-old naïve great tit fledglings ( Parus major ) did not respond differently to a model of a perched predator than to a similarly sized model of a non-predator. Although chicks showed distress responses such as warning calls and freezing behaviour, they did not differentiate between the stimuli. In contrast, wild-caught first-year birds (4 mo old) and adults responded differentially to the two stimuli. Lack of recognition of a perched predator might be one explanation for the high mortality rate found in newly fledged great tits. Our results imply that parental care is not only important for food provisioning, but also to reduce predation risk during the time when fledglings are most vulnerable.     

Females that experience threat are better teachers

Superb fairy-wren (Malurus cyaneus) females use an incubation call to teach their embryos a vocal password to solicit parental feeding care after hatching. We previously showed that high call rate by the female was correlated with high call similarity in fairy-wren chicks, but not in cuckoo chicks, and that parent birds more often fed chicks with high call similarity. Hosts should be selected to increase their defence behaviour when the risk of brood parasitism is highest, such as when cuckoos are present in the area. Therefore, we experimentally test whether hosts increase call rate to embryos in the presence of a singing Horsfield''s bronze-cuckoo (Chalcites basalis). Female fairy-wrens increased incubation call rate when we experimentally broadcast cuckoo song near the nest. Embryos had higher call similarity when females had higher incubation call rate. We interpret the findings of increased call rate as increased teaching effort in response to a signal of threat.     

How does an increase in minimum daily temperatures during incubation influence reproduction in the great tit Parus major?

Temperature variation affects all life stages of organisms, especially early development, and considering global warming, it is urgent to understand precisely its consequences. In egg‐laying species, incubation behaviour can buffer embryo developmental temperature variation and influence offspring development. We experimentally investigated the effect of an increase in minimum daily nest temperature during incubation in the great tit Parus major, by placing a hand warming pad under the nest in the evenings. As compared to controls, the experimental treatment increased nest temperature at night by an average of 4°C, and this increase carried over to the following day. We measured the consequences of this mainly nocturnal temperature increase during incubation on 1) parental behaviour (incubation and nestling feeding), 2) parental health (quantified by body condition, immune status, physiological and oxidative stress) and 3) reproductive success (nestling body condition, growth, i.e. mass gain, hatching and fledging success, and nestling immune status, physiological and oxidative stress). This study yielded three major results. First, we found that heating the nest did not change the duration of incubation as compared to controls. Second, increasing nest temperature during incubation decreased nestling feeding behaviour but did not affect parental health in terms of body condition, immune status, physiological and oxidative stress. Third, nestling mass at hatching was greater but nestling mass gain was slower in heated nests than in control nests, resulting in similar fledging mass. The present study demonstrates that increased environmental temperatures during incubation influenced nestling development in the great tit and especially hatchling mass, which might produce long‐term life history consequences.     

Density dependent effects between three competitive bird species

Summary Density and breeding success of the great tit Parus major, blue tit Parus caeruleus and collared flycatcher Ficedula albicollis were studied in nest box colony in oak forest over a period of 19 years.Intraspecific density dependent clutch size reduction was found with blue tit and great tit. In interspecific relation the high density of blue tits reduced the clutch size of great tits.In the hatching period neither intraspecific nor interspecific density dependence were showed between the tits when the third competitive species, collared flycatcher was present. The collared flycatcher significantly reduced the hatching success of both tit species and the number of fledglings of great tit.The effects of the great tits and combined density of the great and blue tits on the hatching failure and number of fledglings of collared flycatcher were found when the density of the tits was high. There were not significant relationships to the single density of blue tits.The temporal variability of the competition of the three bird species is discussed.     

Sexual imprinting and the origin of obligate brood parasitism in birds

We discuss two pathways along which obligate brood parasitism (OBP) may evolve and examine some of the critical steps that must be passed by letting great tits Parus major be reared by blue tits Parus caeruleus in a field experiment. The cross-fostered chicks survived well in blue tit nests, but their local recruitment and reproductive success was much lower than that of controls. The effect was strongest when great tits grew up with siblings of the host species rather than with conspecific siblings in blue tit nests. The low success seemed to be caused by misimprinting because the cross-fostered birds behaved like blue tits in several aspects (species association, alarm calls, and aggressive response by resident females to caged intruders). Some birds of both sexes were apparently so strongly imprinted that they did not attract or accept a social mate of their own species. We conclude that imprinting may be necessary for OBP to evolve in birds because the parasite must be attracted to the nests of the host species to add eggs and thereby continue the parasitic life cycle. However, strong imprinting may also prevent OBP from occurring if parasitic offspring seek a mate from the host species.     

Incubation prior to clutch completion accelerates embryonic development and so hatch date for eggs laid earlier in a clutch in the great tit Parus major

The ability of parents to respond to changes in food supply within a season will have a large effect on fitness through the number and quality of chicks fledged. Great tits, Parus major, attempt to synchronise their production of chicks with a seasonal food peak, but when food supply fails, hatching asynchrony of chicks provides a mechanism by which some young can be fledged because more developed chicks outcompete their less developed siblings for the reduced parental food supply. We tested whether female great tits can potentially control the degree of hatching asynchrony by using incubation before clutch completion, so that early laid eggs develop faster and hatch sooner. The temperature of an artificial egg placed in 29 nests during the laying period was measured with data loggers, and nocturnal incubation of eggs similar to incubation post clutch completion was recorded in all nests. We then demonstrated that eggs removed from the nest for 72 hour periods prior to clutch completion hatched later than eggs remaining in the nest for the entirety of the laying period. Our results show that variable pre clutch completion incubation (which was mostly nocturnal) can lead to faster embryo development and earlier hatching, so potentially providing a mechanism for adaptive female control of degree of hatching asynchrony.     

Effects of ectoparasites of young on parents' behaviour in a Mediterranean population of Blue Tits

In altricial birds, adults are expected to alleviate the impact of parasitism on their young by increasing parental care. We experimentally investigated the effects of blowfly parasitism on adult behaviour in the nest cavity in a population of Blue Tits in which nestlings are heavily parasitized by blowfly larvae. The behaviour of males and females was different, since only the females carried out nest sanitation activities related to the presence of parasites. The time females devoted to nest sanitation was higher in parasitized nests than in experimentally deparasitized ones, suggesting that chicks and adults might share the costs of blowfly parasitism.     

Corticosterone levels in host and parasite nestlings: Is brood parasitism a hormonal stressor?

Parasite chicks from non-evictor species usually try to monopolize host parental care, thereby increasing considerably the level of food competition in the nest. Here, we propose that brood parasitism is an important stressor for host and parasite nestlings and explore this hypothesis in the non-evictor great spotted cuckoo (Clamator glandarius) and its main hosts, the same-sized black-billed magpie (Pica pica) and the larger carrion crow (Corvus corone). We experimentally created 3-nestling broods of different brood compositions (only cuckoo chicks, only host chicks, or cuckoo and host chicks together) and measured baseline corticosterone levels of nestlings along their developmental period (early, middle and late). We found that brood parasitism increased corticosterone levels in magpie nestlings in the mid and late nestling period compared to those raised in unparasitized nests. Interestingly, carrion crow nestlings from parasitized nests only increased their corticosterone levels in the mid nestling period, when the competition for food with the cuckoo nestling was highest. Our results suggest that brood parasitism could be a potential physiological stressor for host nestlings, especially during the developmental stages where food requirements are highest. Conversely, cuckoo nestlings could be physiologically adapted to high competition levels since they did not show significant differences in corticosterone levels in relation to brood composition.     

Bird predation by tawny owls (Strix aluco L.) and its effect on the reproductive performance of tits

The density of great tit Parus major L. and blue tit Parus caeruleus L. was artificially increased by placing nest-box colonies for these species in the vicinity of the nests of breeding tawny owls during 1993–1997. Bird prey composition in the owl nests, the proportion of parents disappearing from the breeding tit populations and the reproductive performance of the widowed parents were analysed. The frequency of predation on tits by tawny owls was greater in areas where tit density had been artificially increased. Owls preyed more on tits during the feeding period of owlets than during the incubation period and more in years when snow covered the ground during the incubation period than when it did not. Mortality due to predation was male biased and more females lost their mates in populations breeding near tawny owl nests. Reproductive performance of the widowed parents was lower and their body weights were lighter at the end of the nestling period than those found in birds rearing youngs with their mates. Predation by owls increased the between-year turnover in the breeding tit population: widowed parents did not return to the nesting site for the next breeding season.     

Nest biology of urban populations of cavity-nesting birds

Nesting features of four species of cavity-nesting birds—common starling (Sturnus vulgaris L.), great tit (Parus major L.), common redstart (Phoenicurus phoenicurus L.), and pied flycatcher (Ficedula hypoleuca Pall.)—have been studied in city. Under urban conditions, number of eggs in a nest of a redstart is significantly larger, while for great tit and pied flycatcher it is significantly smaller; differences between the populations of starling are insignificant. Success of reproduction for the cavity-nesters is significantly higher in a city and exceeds the reference value for all the species by 10–20%. The number of chicks per a reproduction attempt is also higher in city for all the species. Starling population in city exceeds the reference population in the success of reproduction mainly due to a decrease in the percentage of abandoned eggs. The main reasons for an increase in the reproduction success of other species at the urbanized locality are weakened pressure of predators and a decrease in the percentage of abandoned broods.     

Prolactin and parental behavior in Adélie penguins: effects of absence from nest, incubation length, and nest failure

Adélie penguin (Pygoscelis adeliae) males and females, nesting in Antarctica, alternate attendance at the nest with absences of many days to forage at sea. We investigated the importance of tactile input from egg and chicks on prolactin levels by observing nest attendance patterns and obtaining blood samples (1) during the first nest exchange of the incubation stage, (2) from birds whose incubation period was artificially increased or decreased by about 10 days, and (3) from birds whose nests had failed. Prolactin levels in females after 8 to 11 days of absence from the breeding colony did not differ from those in incubating males and did not change after females resumed incubation. Moving eggs between nests resulted in nests in which chicks hatched after about 26, 36 (normal), or 46 days. Duration of incubation did not affect prolactin levels in the parents measured during incubation, at the pip stage, hatch stage, or early brood stage. Adults first left their chicks unguarded on about the same calendar date, regardless of chick age. However, chicks from long incubation nests averaged 8 days younger when they were left unguarded than chicks from control or short-incubation nests. In females, there was no effect of nest failure on prolactin levels. In males, prolactin levels were slightly lower after nest failure than in males tending nests. Testosterone was significantly higher in males after nest failure than in males still tending nests. Prolactin is elevated in Adélie penguins as part of the program of cyclical hormonal changes that accompany the lengthy reproductive season and is relatively independent of tactile input. Sustained prolactin secretion is probably required for the maintenance of parental behavior in offshore feeding species that must be absent from the nest for many days at a time.     

Individual benefits of nestling begging: experimental evidence for an immediate effect, but no evidence for a delayed effect

The evolutionary stability of honest signalling by offspring is thought to require that begging displays be costly, so the costs and benefits of begging--and whether they are experienced individually or by the whole brood--are crucial to understanding the evolution of begging behaviour. Begging is known to have immediate individual benefits (parents distribute more food to intensely begging individuals) and delayed brood benefits (parents increase provisioning rate to the brood), but the possibility of delayed individual benefits (previous begging affects the current distribution of food) has rarely, if ever, been researched. We did this using playback of great tit Parus major chick begging and a control sound from either side of the nest. Male parents fed chicks close to the speaker more when great tit chick begging, but not other stimuli, was played back. In contrast, there was no effect of playback at the previous visit on the chicks that male parents fed. We have thus demonstrated an immediate individual benefit to begging, but found no evidence of a delayed individual benefit in this species.     

Parasites, predators and the duration of developmental periods

Nest predation has been suggested to be the most important environmental factor determining the evolution of incubation and nestling periods in birds. This paradigm is based on a number of comparative analyses that have shown correlations between rates of nest predation and the relative duration of the nestling period. A second important natural selection pressure on reproduction is caused by parasitism. Here, I investigated the independent effects of daily nest predation rates and daily nest mortality rates due to parasites to assess the relative importance of these two environmental factors as determinants of the relative duration of incubation and nestling periods in birds, using the 43 species for which information on parasite-induced nestling mortality was available. The relative duration of the incubation period was negatively related to daily mortality rate due to parasitism, while nest predation had no significant effect. The relative duration of the nestling period was not significantly related to daily rate of mortality due to parasitism or nest predation. Daily nest predation rate was significantly negatively correlated with daily mortality rate due to parasitism in the subsample of species, for which some mortality due to parasitism was recorded. These findings are mainly based on hole nesters and colonially breeding species, because estimates of parasite-induced mortality generally is unavailable from open nesting and solitarily breeding species, precluding any generalizations to these species. However, the conclusions did not change after statistically controlling for nest site and sociality. These results suggest that developmental rates may be more influenced by the effects of parasites rather than nest predators, and that parasites constitute a neglected selective force affecting developmental rates.     

Shared parental care is costly for nestlings of common cuckoos and their great reed warbler hosts

Obligate avian brood parasitism typically involves one of 2strategies: parasite chicks are either 1) virulent and evictall other eggs and nest mates to be raised alone or 2) moretolerant and share foster parental care with host chicks forsome or the entirety of the nestling period. We studied theconsequences of experimentally forced mixed broods of age-matchedone common cuckoo (Cuculus canorus) and 2 great reed warbler(Acrocephalus arundinaceus) chicks. In these broods, both cuckooand host chicks grew slower than did either individual cuckoosor great reed warblers in broods of 1 parasite or 3 host chicks,respectively. Video records showed that in mixed broods, cuckoochicks received feedings less frequently than the 33% predictedby chance at 4 days of age but parental food allocations increasedto chance levels at 8 days of age. The consistent patterns oflower growth rates arose even though chicks in broods of 1 parasiteand 2 hosts received the largest prey items per feeding. Inaddition, several other measures of parental provisioning alsodid not predict species and brood-specific differences in nestlinggrowth rates across the different treatments. However, variationin begging displays and its specific costs on host and parasitechicks in the different nest treatments were not quantifiedin this study. We conclude that young of nest mate–evictorcommon cuckoos benefit from the sole occupancy of host nestsin part owing to an initial competitive disadvantage for parentalcare in broods with age-matched great reed warbler chicks.     

The Evolution of Intraspecific Brood Parasitism in Birds and Insects

Many species of birds and insects engage in intraspecific brood parasitism (IBP), when a female lays eggs in the nest of a conspecific and leaves without providing parental care. These visiting females may also act to cooperate with a primary female, staying to provide parental care. Therefore, IBP and cooperative breeding can be considered extremes on a continuum of parental care provided by a secondary female. When a secondary female abandons a nest, she creates an asymmetry in parental care between herself and the host. While models of asymmetry in reproductive allocation have focused directly on relatedness between females, we lack an appropriate theoretical framework that addresses the effects of relatedness on parental care asymmetry. Here, I present an evolutionarily stable strategy (ESS) model that predicts the conditions under which IBP is favored over cooperation and solitary breeding. Intraspecific brood parasitism is less likely to evolve (relative to cooperation and solitary breeding) as the relatedness between a host and parasite increases. It can evolve, however, if parasites achieve a high overall fecundity relative to solitary females. Constraints on solitary breeding can further promote IBP under some circumstances. Cooperation is favored when relatedness is high and reproductive skew is low. This model makes several predictions regarding the conditions under which IBP may evolve, motivating a variety of experimental approaches.     

Parents, predators, parasites, and the evolution of eggshell colour in open nesting birds

The colourful surface of birds’ eggshells varies dramatically between species, but the selective pressures driving this variation remain poorly understood. We used a large comparative dataset to test several hypotheses proposed to explain the evolution of eggshell colouration. We tested the hypothesis that predation pressure might select for cryptic eggshells by examining the relationship between predation rate and egg colouration. We found that predation rates were positively related to eggshell brightness. The blackmail hypothesis suggests that females lay colourful eggshells to coerce males into providing additional care during incubation to keep colourful eggs covered. According to this hypothesis, conspicuous eggs should be found in situations with high risk of visual detection from predators or brood parasites. In support of this hypothesis, proportional blue-green chroma was positively related to parasitism risk, and eggs with higher proportional blue-green chroma or higher ultraviolet chroma received higher combined parental nest attendance during the incubation period. The sexual signalling hypothesis states that blue-green colour indicates female quality; however, we did not find that blue-green eggshell colour was greater in species where males participate in any form of parental care, and relative male provisioning was unrelated to blue-green eggshell chroma. We found some support for the hypothesis that brood parasitism may select for high inter-clutch variation in eggshell colour to facilitate egg recognition. In our dataset, parasitism risk was negatively related to inter-clutch repeatability of blue-green chroma. Our study highlights the diversity of selection pressures acting on the evolution of eggshell colour in birds and provides suggestions for novel areas of future key research direction.     

Sympatric and allopatric combinations of hen fleas and great tits: a test of the local adaptation hypothesis

To test whether allopatric nest parasites differ from sympatric ones in their effect on various life history traits of their avian host, I designed a cross-transfer experiment in which hen fleas (Ceratophyllus gallinae) were exchanged between great tit (Parus major) nests in two geographically widely separated areas. In neither of the areas did allopatric fleas influence body mass, tarsus length, wing length, duration of nestling period or mortality in great tit nestlings more severely than did sympatric fleas. Duration of incubation was also similar among females independent of experimental treatment. This lack of difference between allopatric and sympatric fleas is hypothesized to reflect the comparative harmlessness of hen fleas for their hosts.     

Darwin's Finch Begging Intensity Does Not Honestly Signal Need in Parasitised Nests

Parental care should be selected to respond to honest cues that increase offspring survival. When offspring are parasitised, the parental food compensation hypothesis predicts that parents can provision extra food to compensate for energy loss due to parasitism. Chick begging behaviour is a possible mechanism to solicit increased feeding from attending parents. We experimentally manipulated parasite intensity from Philornis downsi in nests of Darwin's small ground finch (Geospiza fuliginosa) to test its effects on chick begging intensity and parental food provisioning. We used in‐nest video recordings of individually marked chicks to quantify nocturnal parasite feeding on chicks, subsequent diurnal chick begging intensity and parental feeding care. Our video analysis showed that one chick per brood had the highest parasite intensity during the night (supporting the tasty chick hypothesis) and weakest begging intensity during the day, which correlated with low parental care and rapid death. We observed sequential chick death on different days rather than total brood loss on a given day. Our within‐nest video images showed that (1) high nocturnal larval feeding correlated with low diurnal begging intensity and (2) parent birds ignored weakly begging chicks and provisioned strongly begging chicks. Excluding predation, all parasite‐free chicks survived (100% survival) and all parasitised chicks died in the nest (100% mortality). Weak begging intensity in parasitised chicks, which honestly signalled recent parasite attack, was not used as a cue for parental provisioning. Parents consistently responded to the strongest chick in both parasitised and parasite‐free nests.     

Constraints on host choice: why do parasitic birds rarely exploit some common potential hosts?

1. Why are some common and apparently suitable resources avoided by potential users? This interesting ecological and evolutionary conundrum is vividly illustrated by obligate brood parasites. Parasitic birds lay their eggs into nests of a wide range of host species, including many rare ones, but do not parasitize some commonly co-occurring potential hosts. 2. Attempts to explain the absence of parasitism in common potential hosts are limited and typically focused on single-factor explanations while ignoring other potential factors. We tested why thrushes Turdus spp. are extremely rarely parasitized by common cuckoos Cuculus canorus despite breeding commonly in sympatry and building the most conspicuous nests among forest-breeding passerines. 3. No single examined factor explained cuckoo avoidance of thrushes. Life-history traits of all six European thrush species and the 10 most frequently used cuckoo hosts in Europe were similar except body/egg size, nest design and nestling diet. 4. Experiments (n = 1211) in several populations across Europe showed that host defences at egg-laying and incubation stages did not account for the lack of cuckoo parasitism in thrushes. However, cross-fostering experiments disclosed that various factors during the nestling period prevent cuckoos from successfully parasitizing thrushes. Specifically, in some thrush species, the nest cup design forced cuckoo chicks to compete with host chicks with fatal consequences for the parasite. Other species were reluctant to care even for lone cuckoo chicks. 5. Importantly, in an apparently phylogenetically homogenous group of hosts, there were interspecific differences in factors responsible for the absence of cuckoo parasitism. 6. This study highlights the importance of considering multiple potential factors and their interactions for understanding absence of parasitism in potential hosts of parasitic birds. In the present study, comparative and experimental procedures are integrated, which represent a novel approach that should prove useful for the understanding of interspecific ecological relationships in general.