The nest defence by the red‐backed shrike Lanius collurio – support for the vulnerability hypothesis

The majority of altricial bird species defend their brood against predators more intensively in nestlings rather than eggs stage. Several hypotheses have been proposed to explain this difference. The majority of existing experimental studies have recorded a gradually increasing intensity of nest defence supporting the reproductive value hypothesis. We have compared nest defence in two nesting stages of the red‐backed shrike against two predators of adult birds and against two predators of nests. While the nests with nestlings were defended by parents against three out of four predators, nests with eggs were almost not defended at all. This rapid change in parent nest defence supports rather the vulnerability hypothesis, predicting that the threat to nests with nestlings increases rapidly after hatching, as they became more conspicuous due to their begging and parental provisioning. Unlike most of the species tested previously, the red‐backed shrike uses very vigorous mobbing towards predators. We suggest that the occurrence of this active mobbing (strikes, including physical contact) is a good proxy of the current threat to the nest.     

Contributions of feather microstructure to eider down insulation properties

Insulation is an essential component of nest structure that helps provide incubation requirements for birds. Many species of waterfowl breed in high latitudes where rapid heat loss can necessitate a high energetic input from parents and use down feathers to line their nests. Common eider Somateria mollissima nest down has exceptional insulating properties but the microstructural mechanisms behind the feather properties have not been thoroughly examined. Here, we hypothesized that insulating properties of nest down are correlated to down feather (plumule) microstructure. We tested the thermal efficiency (fill power) and cohesion of plumules from nests of two Icelandic colonies of wild common eiders and compared them to properties of plumules of wild greylag goose Anser anser. We then used electron microscopy to examine the morphological basis of feather insulating properties. We found that greylag goose down has higher fill power (i.e. traps more air) but much lower cohesion (i.e. less prone to stick together) compared to common eider down. These differences were related to interspecific variation in feather microstructure. Down cohesion increased with the number of barbule microstructures (prongs) that create strong points of contact among feathers. Eider down feathers also had longer barbules than greylag goose down feathers, likely increasing their air‐trapping capacity. Feather properties of these two species might reflect the demands of their contrasting evolutionary history. In greylag goose, a temperate, terrestrial species, plumule microstructure may optimize heat trapping. In common eiders, a diving duck that nests in arctic and subarctic waters, plumule structure may have evolved to maximize cohesion over thermal insulation, which would both reduce buoyancy during their foraging dives and enable nest down to withstand strong arctic winds.     

The effects of hatching asynchrony on growth and mortality patterns in Eurasian Hoopoe Upupa epops nestlings

Growth is a fundamental life history trait in all organisms and is closely related to individual fitness. In altricial birds, growth of many traits is restricted to the short period between hatching and fledging and strongly depends on the amount of food that parents deliver and the extent of hatching asynchrony. However, empirical studies of energy allocation to growth of different body size traits as a function of hatching asynchrony are scarce. We studied growth and mortality of Eurasian Hoopoe Upupa epops, a species with a long breeding season and high brood size variance, whose nestlings show pronounced hatching asynchrony, in order to test how hatching asynchrony affects different growth traits in the context of territory quality, season and brood size. The growth of five body traits (body mass, and lengths of tarsus, third primary, bill and longest crest feather) was investigated to understand how it was affected by brood size, hatching date and order, and territory quality. In total, 241 nestlings from 39 nests were measured every 4 days in 2014 in south‐western Switzerland. Brood size, hatching date and hatching order had the strongest influence on growth trajectories, although tarsus growth was only marginally affected by these variables. Nestlings that hatched earlier than their siblings were heavier and had longer third primaries, bills and crest feathers compared with later‐hatched siblings. In territories of high quality, hatching order differences disappeared for body mass growth, but persisted for lengths of third primary, bill and crest feathers. Brood size was inversely associated with third primary, bill and crest feather lengths, but positively associated with body mass. Nestling mortality was higher in later‐hatched nestlings and in broods that were raised in territories of lower quality. Our study shows that in nestlings, energy was allocated differentially between body traits and this allocation interacted with hatching order and territory quality. Rapid mass gain by nestlings was prioritized in order to increase competitive ability. Our results provide support for the brood reduction hypothesis as an explanation of hatching asynchrony in Hoopoes.     

Environmental conditions but not nest composition affect reproductive success in an urban bird

Adjusting the composition of their nests, breeding birds can influence the environmental conditions that eggs and offspring experience. Birds often use feathers to build nests, presumably due to their insulating properties. The amount of feathers in nests is often associated with increased nestling survival and body condition. However, it is unclear whether these putative beneficial effects of adding feathers to nests are relevant in a wide range of environmental conditions. Here, we combine data on weather conditions and feathers in nests (i.e., nest composition) to investigate their relative contribution to reproductive success in the Eurasian tree sparrow (Passer montanus). Specifically, we investigate whether the effect of weather conditions on breeding success is modulated by the amount of feathers added to the nest. We found a strong negative effect of rainfall on the number of nestlings that successfully fledged per breeding attempt, but this negative effect was not mitigated by the amount of feathers in nests. We also found that the amount of feathers in nests varied along the breeding season, with nests containing more feathers early in the breeding season, when temperatures were lower. Despite considerable variation in nest composition, our results do not suggest an important role of feathers in nests protecting eggs or nestling tree sparrows against fluctuations in environmental conditions.     

Colour composition of nest lining feathers affects hatching success of barn swallows,Hirundo rustica (Passeriformes: Hirundinidae)

Many bird species use feathers as lining material, and its functionality has traditionally been linked to nest insulation. However, nest lining feathers may also influence nest detection by predators, differentially affect reproductive investment of mates in a post‐mating sexual selection process, and affect the bacterial community of the nest environment. Most of these functions of nest lining feathers could affect hatching success, but the effect might vary depending on feather coloration (i.e. pigmented versus white feathers). This would be the case if coloration is related to: (1) thermoregulatory properties; (2) attractiveness of feathers in the nest for mates; (3) eggshell bacterial density. All of these hypothetical scenarios predict that feathers of different colours would differentially affect the hatching success of birds, and that birds should preferentially choose the most beneficial feather colour for lining their nests. Results from two different experiments performed with a population of Danish barn swallow, Hirundo rustica, were in accordance with these predictions. First, H. rustica preferentially selected white experimentally offered feathers for lining their nests. Second, the experimental manipulation of the feather colour composition of nests of H. rustica had a significant effect on hatching success. Experimental nests with more white feathers added at the beginning of incubation had a lower probability of hatching failures, suggesting differential beneficial effects of lining nests with feathers of this colour. We discuss the relative importance of hypothetical functional scenarios that predicted the detected associations, including those related to sexual selection or to the community of microorganisms associated with feathers of different colours. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 67–74.     

The effects of nest size and insulation on thermal properties of tree swallow nests

The nest environment can have important influences on incubation behavior and nestling development in birds. Nest thermal properties, particularly nest composition and size, can have a major influence on heat loss. To examine the role of nest size and insulation on clutch cooling rates, we collected tree swallow Tachycineta bicolor nests and measured the cooling rate of eggs in a controlled thermal environment. We also examined the thermal benefits of nest feathers by comparing the cooling rates of nests with and without feathers. Nests with more feather insulation and heavier, deeper cupped nests cooled at slower rates. In addition, nests with feathers cooled at much slower rates than did the same nests without feathers. Our results show that nest insulation and size play important roles in nest cooling rates, which may ultimately affect incubation costs and thus reproductive performance.     

MAGPIE HOST MANIPULATION BY GREAT SPOTTED CUCKOOS: EVIDENCE FOR AN AVIAN MAFIA?

Why should the hosts of brood parasites accept and raise parasitic offspring that differ dramatically in appearance from their own? There are two solutions to this evolutionary enigma. (1) Hosts may not yet have evolved the capability to discriminate against the parasite, or (2) parasite-host systems have reached an evolutionary equilibrium. Avian brood parasites may either gain renesting opportunities or force their hosts to raise parasitic offspring by destroying or preying upon host eggs or nestlings following host ejection of parasite offspring. These hypotheses may explain why hosts do not remove parasite offspring because only then will hosts avoid clutch destruction by the cuckoo. Here we show experimentally that if the egg of the parasitic great spotted cuckoo Clamator glandarius is removed from nests of its magpie Pica pica host, nests suffer significantly higher predation rates than control nests in which parasite eggs have not been removed. Using plasticine model eggs resembling those of magpies and observations of parasites, we also confirm that great spotted cuckoos that have laid an ejected egg are indeed responsible for destruction of magpie nests with experimentally ejected parasite eggs. Cuckoos benefit from destroying host offspring because they thereby induce some magpies to renest and subsequently accept a cuckoo egg.     

Is Carrying Feathers a Sexually Selected Trait in House Sparrows?

Males in several avian groups carry specialised materials as part of their courtship display. Females may vary their investment in reproduction in relation to their mates' attractiveness. The house sparrow (Passer domesticus) population on Dassen Island, South Africa, builds domed nests in the branches of trees. These nests are more or less spherical structures with a deep cup lined largely with feathers. When males collect feathers, they call to females and display the feather before it is added to the nest. We examined whether carrying feathers provides females with an index of male quality, which correlates with their subsequent investment in parental care. Additionally, we studied the potential importance of feathers in nest insulation. Feathers were collected mainly by males. Males also brought larger feathers, and deposited more feathers in nests, than females. Number of trips with feathers – which increased after feathers were experimentally removed from nests – and number of feathers brought varied among males. Volume of feathers influenced females' investment in reproduction and positively correlated with clutch size and chick feeding rates. We found more feathers during incubation and immediately after chicks hatched, when nest heat requirements peak. Furthermore, number of trips with feathers and number and size of feathers were greater during incubation. Our results suggest that this feather‐carrying behaviour by male house sparrows influences maternal reproduction investment and could be a kind of parental care per se by supplementing nest insulation.     

Brood size manipulations reveal relationships among physiological performance,body condition and plumage colour in Northern Flicker Colaptes auratus nestlings

Visual signals of quality in offspring, such as plumage colour, should honestly advertise need and/or body condition, but links between nutritional status, physiological performance and the expression of colours are complex and poorly understood. We assess how food stress during rearing affected two physiological measures (T‐cell‐mediated immune function and corticosterone level in feathers: CORT_f) and how these two variables were related to carotenoid and melanin coloration in Northern Flicker Colaptes auratus nestlings. We were also interested in how these two physiological measures were influenced by the sex of the nestling. We experimentally manipulated brood size to alter levels of food availability to nestlings during development. We measured carotenoid‐based colour (chroma and brightness) in wing feathers and the size of melanin spots on breast feathers. In agreement with our prediction, nestlings in the reduced brood treatment had better body condition and stronger immune responses than those in the control and brood enlargement treatments. This supports the hypothesis that immune responses are energetically costly. In contrast, CORT_f was not related to nestling body condition or sex and was unaffected by brood size manipulation. Nestlings of both sexes with stronger T‐cell‐mediated immune responses had larger melanin spots but only males with higher immune responses also had brighter flight feathers. Feather brightness decreased with increasing CORT_f levels. Our study is one of the few to examine the relationship between multiple physiological and plumage measures in nestlings and shows that plumage colour and immune function signalled body condition of nestlings, but that feather corticosterone levels did not.     

Discrimination and ejection of eggs and nestlings by the fan-tailed gerygone from New Caledonia

Nestling rejection is a rare type of host defense against brood parasitism compared with egg rejection. Theoretically, host defenses at both egg and nestling stages could be based on similar underlying discrimination mechanisms but, due to the rarity of nestling rejector hosts, few studies have actually tested this hypothesis. We investigated egg and nestling discrimination by the fan-tailed gerygone Gerygone flavolateralis, a host that seemingly accepts nonmimetic eggs of its parasite, the shining bronze-cuckoo Chalcites lucidus, but ejects mimetic parasite nestlings. We introduced artificial eggs or nestlings and foreign gerygone nestlings in gerygone nests and compared begging calls of parasite and host nestlings. We found that the gerygone ejected artificial eggs only if their size was smaller than the parasite or host eggs. Ejection of artificial nestlings did not depend on whether their color matched that of the brood. The frequency of ejection increased during the course of the breeding season mirroring the increase in ejection frequency of parasite nestlings by the host. Cross-fostered gerygone nestlings were frequently ejected when lacking natal down and when introduced in the nest before hatching of the foster brood, but only occasionally when they did not match the color of the foster brood. Begging calls differed significantly between parasite and host nestlings throughout the nestling period. Our results suggest that the fan-tailed gerygone accepts eggs within the size range of gerygone and cuckoo eggs and that nestling discrimination is based on auditory and visual cues other than skin color. This highlights the importance of using a combined approach to study discrimination mechanisms of hosts.     

Co‐infections by malaria parasites decrease feather growth but not feather quality in house martin

During moult, stressors such as malaria and related haemosporidian parasites (e.g. Plasmodium and Haemoproteus) could affect the growth rate and quality of feathers, which in turn may compromise future reproduction and survival. Recent advances in molecular methods to study parasites have revealed that co‐infections with multiple parasites are frequent in bird–malaria parasite systems. However, there is no study of the consequences of co‐infections on the moult of birds. In house martins Delichon urbica captured and studied at a breeding site in Europe during 11 yr, we measured the quality and the growth rate of tail feathers moulted in the African winter quarters in parallel with the infection status of blood parasites that are also transmitted on the wintering ground. Here we tested if the infection with two haemosporidian parasite lineages has more negative effects than a single lineage infection. We found that birds with haemosporidian infection had lower body condition. We also found that birds co‐infected with two haemosporidian lineages had the lowest inferred growth rate of their tail feathers as compared with uninfected and single infected individuals, but co‐infections had no effect on feather quality. In addition, feather quality was negatively correlated with feather growth rate, suggesting that these two traits are traded‐off against each other. We encourage the study of haemosporidian parasite infection as potential mechanism driving this trade‐off in wild populations of birds.     

Behavioural responses to ectoparasites in pied flycatchers Ficedula hypoleuca: an experimental study

Nests of cavity‐nesting birds usually harbor some species of haematophagous ectoparasites that feed on the incubating adults and nestlings. Given the negative impact of ectoparasites on nestlings there will be selection on hosts to reduce parasite infestations through behavioural means. We have experimentally reduced the abundance of all ectoparasites in nests of pied flycatchers Ficedula hypoleuca to explore both whether there are changes in the frequency and duration of putative anti‐parasite behaviours by tending adults, as well as whether such anti‐parasite behaviours are able to compensate for the deleterious effects that parasites may have on nestlings. Heat treatment of nests substantially decreased the density of ectoparasites, and thereby positively affected nestling growth. The frequency and intensity of female grooming and nest sanitation behaviours during the incubation and nestling periods decreased as a consequence of the experimental reduction of ectoparasite infestation. Although nestlings begged more intensely in infested nests, the experiment had no significant effect on parental provisioning effort. Reduction of parasites resulted in larger nestlings shortly before fledging and increased fledging success. This study shows a clear effect of a complete natural nest ectoparasite fauna on parental behaviour at the nest and nestling growth in a cavity‐nesting bird. Although ectoparasites induce anti‐parasite behaviours in females, these behaviours are not able to fully remove parasite's deleterious effects on nestling growth and survival.     

Intraseasonal variation in immune defence,body mass and hematocrit in adult house martins Delichon urbica

The intensity of parasite infections often increases during the reproductive season of the host as a result of parasite reproduction, increased parasite transmission and increased host susceptibility. We report within‐individual variation in immune parameters, hematocrit and body mass in adult house martins Delichon urbica rearing nestlings in nests experimentally infested with house martin bugs Oeciacus hirundinis and birds rearing nestlings in initially parasite‐free nests. From first to second broods body mass and hematocrit of breeding adult house martins decreased. In contrast leucocytes and immunoglobulins became more abundant. When their nests were infested with ectoparasites adults lost more weight compared with birds raising nestlings in nests treated with pyrethrin, whereas the decrease in hematocrit was more pronounced during infection with blood parasites. Neither experimental infestation with house martin bugs nor blood parasites had a significant effect on the amount of immune defences.     

Supplementary feeding increases nestling feather corticosterone early in the breeding season in house sparrows

Several studies on birds have proposed that a lack of invertebrate prey in urbanized areas could be the main cause for generally lower levels of breeding success compared to rural habitats. Previous work on house sparrows Passer domesticus found that supplemental feeding in urbanized areas increased breeding success but did not contribute to population growth. Here, we hypothesize that supplementary feeding allows house sparrows to achieve higher breeding success but at the cost of lower nestling quality. As abundant food supplies may permit both high‐ and low‐quality nestlings to survive, we also predict that within‐brood variation in proxies of nestling quality would be larger for supplemental food broods than for unfed broods. As proxies of nestling quality, we considered feather corticosterone (CORT_f), body condition (scaled mass index, SMI), and tarsus‐based fluctuating asymmetry (FA). Our hypothesis was only partially supported as we did not find an overall effect of food supplementation on FA or SMI. Rather, food supplementation affected nestling phenotype only early in the breeding season in terms of elevated CORT_f levels and a tendency for more variable within‐brood CORT_f and FA. Early food supplemented nests therefore seemed to include at least some nestlings that faced increased stressors during development, possibly due to harsher environmental (e.g., related to food and temperature) conditions early in the breeding season that would increase sibling competition, especially in larger broods. The fact that CORT_f was positively, rather than inversely, related to nestling SMI further suggests that factors influencing CORT_f and SMI are likely operating over different periods or, alternatively, that nestlings in good nutritional condition also invest in high‐quality feathers.     

Is the structural and psittacofulvin‐based coloration of wild burrowing parrots Cyanoliseus patagonus condition dependent?

The bright colours of parrots are caused by psittacofulvin pigments, which appear unique to this Order, and by structural colours. We measured red (psittacofulvin), green (mixed) and blue (structural) colours of wild burrowing parrots Cyanoliseus patagonus of northeastern Patagonia, Argentina, and measured nestlings regularly to obtain data on breeding success and nestling growth. As adult feathers are moulted outside the breeding season, adult body condition could not be measured directly during feather growth, and climatic conditions were used as an indirect parameter. The colony of burrowing parrots is surrounded by Monte steppe habitat, the breeding success has been shown to depend strongly on the climatic patterns. The area experienced a drought with very poor breeding success as well as a year of above‐average precipitation during the study period, serving as a natural experiment. We thus analysed the variability of colouration within the population among and within breeding seasons. We observed strong inter‐annual differences in nestling and adult colouration. Nestlings grew blue feathers with lower achromatic brightness during better conditions, and when controlling for year effects, nestlings with higher mass and from more successful families also had blue feathers with lower achromatic brightness. Adult blue feathers showed the same trend, with lower achromatic brightness in the moult following breeding seasons of better conditions. In contrast, during better conditions, adults grew red feathers with higher achromatic brightness and the colour hue was also affected, and the hue of the red region of nestlings varied with the hatching order. The colour of all three regions of nestlings varied between nests, and the colour of the red region of adult males positively correlated with breeding success (clutch size, brood size). In summary, the present data suggest that environmental conditions contribute to variability in both structural and the psittacofulvin‐based colours of wild burrowing parrots.     

Fault bars timing and duration: the power of studying feather fault bars and growth bands together

Growth bands and fault bars, widespread features of feathers that form during regeneration, have largely been studied independently. Growth bands result from normal regeneration: each pair of dark/light bands forms every 24 h. Fault bars are a response to stress during regeneration, creating a translucent line that can break the feather. We studied the relative position and width of these two structures in feathers of nestling and adult white storks Ciconia ciconia. We first confirmed that one growth band represents 24 h of feather regeneration. Fault bars did not occur at random within growth bands: 65.7% (in nestlings) and 45.6% (in adults) of them occurred in one out of six defined segments within a growth band, namely that segment corresponding to the first one‐third of night time hours. The width of fault bars relative to growth bands suggested that fault bars were produced during a median (range) of 7.0 h (2.7–27.0) in nestlings and 3.7 h (1.8–7.9) in adults. Fault bars were concentrated at feather tips in nestlings, but at central locations in adult feathers. Our results suggest that, in general, fault bars are a discrete event of a finite duration occurring mainly during the night (particularly in nestlings). This, along with current knowledge, suggests that acute stressors, rather than chronic ones, are responsible for fault bar formation. Thus, such acute punctual stressors (a matter of minutes) can have long‐lasting (months?years) physiological effects due to the wing load increase from feather breakage caused by fault bars.     

Synchronization of laying by great spotted cuckoos and recognition ability of magpies

Brood parasites rely entirely on the parental care of host species to raise the parasitic nestlings until independence. The reproductive success of avian brood parasites depends on finding host nests at a suitable stage (i.e. during egg laying) for parasitism and weakly defensive (i.e. non‐ejector) hosts. Finding appropriate nests for parasitism may, however, vary depending on ecological conditions, including parasite abundance in the area, which also varies from one year to another and therefore may influence coevolutionary relationships between brood parasites and their hosts. In this scenario, we explored: 1) the degree of laying synchronization between great spotted cuckoos Clamator glandarius and magpies Pica pica during two breeding seasons, which varied in the level of selection pressure due to brood parasitism (i.e. parasitism rate); 2) magpie responses to natural parasitism in the pre‐laying period and successfulness of parasitic eggs laid at this stage; and 3) magpie responses to experimental parasitism performed at different breeding stages. We found that, during the year of higher parasitism rate, there was an increase in the percentage of parasitic eggs laid before magpies started laying. However, the synchronization of laying was poor both years regardless of the differences in the parasitism rate. The ejection rate was significantly higher during the pre‐egg‐laying and the post‐hatching stages than during the laying stage, and hatching success of parasitic eggs laid during the pre‐egg‐laying stage was zero. Thus, non‐synchronized parasitic eggs are wasted and therefore poor synchronization should be penalized by natural selection. We discuss four different hypotheses explaining poor synchronization.     

Philornis infection in blue‐black grassquits: impact on nestlings and risk factors involved

Parasitic botfly larvae (Philornis ssp., Diptera: Muscidae) are found in nests of several bird taxa, although prevalence and impact on nestling survival vary considerably among species. Here we describe patterns of botfly infestation in blue‐black grassquit Volatinia jacarina nestlings. We identified the most typically affected nestling body parts and assessed parasite prevalence, impact on nestling survival, and changes in nestling body shape. Additionally, we tested whether climatic conditions, nest morphology and habitat characteristics are associated with larvae abundance. Blue‐black grassquits had low breeding success (16% of eggs/nestlings survived to fledged; 19% of the nests fledged at least one), but most failures resulted from predation by vertebrate predators. We estimated that 1% of nestlings died due to botfly infestation, and the number of subcutaneous larvae (range 1–18) in a nestling's body did not predict fledging success. Infected chicks exhibited higher tarsus asymmetry. Thus, we argue that although botflies had a small impact on offspring survival, they may reduce fitness in adulthood. There was no evidence that environmental conditions and nest morphology are linked to the number of larvae on nestlings. Nesting areas with higher food supply had lower infestation rates. Possibly, food‐rich habitats allow parents to invest more time in offspring care (brooding nestlings), thus protecting them from fly attacks. Alternatively, vegetation composition could influence local invertebrate diversity, which could provide a natural trophic buffer against adult Philornis. The present study brings to light new perspectives concerning bird–botfly interaction.     

Rejection of brood‐parasitic shiny cowbird Molothrus bonariensis nestlings by the firewood‐gatherer Anumbius annumbi?

Costs imposed by brood parasitic birds exert strong selection on their hosts to avoid parasitism. While egg rejection is a common defence, nestling rejection is rarer and less well understood. Theoretical models suggest that among non‐evicting parasites such as cowbirds nestling rejection can only evolve when levels of parasitism are high. Here we describe a possible case of early rejection of cowbird nestlings, by an infrequently parasitised host, the firewood‐gatherer Anumbius annumbi. Firewood‐gatherers accepted most shiny cowbird Molothrus bonariensis eggs despite clear differences in coloration. Cowbird eggs usually hatched 4–5 d before host eggs. All parasitic nestlings died within 48 h, and hosts continued their breeding attempts. Nestling death was most likely due to neglect since little food was found in the stomach of dead nestlings. Feeding neglect could be due to differences in visual or acoustic appearance between host and parasite hatchlings. Alternatively, hosts may refrain from feeding nestlings that hatch too early compared to their normal incubation time. At the moment our data do not allow distinction between active nestling recognition or cowbird nestling failure due to the unsuitability of the firewood‐gatherer as a host (i.e. too long incubation). Experiments are needed to tease these alternatives apart.