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.     

Incubation Feeding and Nest Attentiveness in a Socially Monogamous Songbird: Role of Feather Colouration,Territory Quality and Ambient Environment

Parental investment and environmental conditions determine reproductive success in wild‐ranging animals. Parental effort during incubation, and consequently factors driving it, has profound consequences for reproductive success in birds. The female nutrition hypothesis states that high male feeding enables the incubating female to spend more time on eggs, which can lead to higher hatching success. Moreover, both male and female parental investment during incubation might be signalled by plumage colouration. To test these hypotheses, we investigated relationships between male and female incubation behaviour and carotenoid and melanin‐based plumage colouration, territory quality and ambient temperature in the Great Tit Parus major. We also studied the effect of female incubation behaviour on hatching success. Intensity of male incubation feeding increased with lower temperatures and was higher in territories with more food supply, but only in poor years with low overall food supply. Female nest attentiveness increased with lower temperatures. Plumage colouration did not predict incubation behaviour of either parent. Thus, incubation behaviour of both parents was related mainly to environmental conditions. Moreover, there was no relationship between male incubation feeding, female nest attentiveness and hatching success. Consequently, our data were not consistent with the female nutrition hypothesis.     

Hissing calls improve survival in incubating female great tits (Parus major)

Nest predation is among the most important selective pressure shaping nest-site selection and nest defense behavior in many avian species. In this study, we tested whether the production of one such nest defense behavior—hissing calls—may improve survival of incubating female great tits (Parus major). We found that 72.5 % of incubating females gave hissing calls when they were exposed to a stuffed woodpecker in their nest boxes. The repeatability of the number of hissing calls given was high, as was the latency to give the call. Additionally, natural nest predators attacked hissing and nonhissing females equally often. However, hissing females survived better than silent females. We tested responses of feral cats to playbacks of hissing call during their attacks of nest boxes and found that hissing calls prevented the predator attacks. Taken together, our findings indicate that hissing calls can deter predator attacks and potentially increase survival rates of nesting great tits or their offspring, or both. The propensity to give hissing calls may be related to personality type of incubating female great tits, which needs to be tested experimentally.     

Interactive vocal communication at the nest by parent Great Tits Parus major

Although most bird species show monogamous pair bonds and bi‐parental care, little is known of how mated birds coordinate their activities. Whether or not partners communicate with each other to adjust their behaviour remains an open question. During incubation and the first days after hatching, one parent – generally the female – stays in the nest for extended periods, and might depend on acoustic communication to exchange information with its mate outside. The Great Tit Parus major is an interesting study system to investigate intra‐pair communication at the nest because males address songs to their mate while she is in the nest cavity, and females answer the male from the cavity with calls. However, the function of this communication remains unknown. In this study, we recorded the vocalizations and observed the resulting behaviour of Great Tit pairs around the nest at different breeding stages (laying, incubation and chick‐rearing). We observed vocal exchanges (vocalization bouts, alternated on the same tempo, between the female inside the nest and her male outside) in three contexts with different outcomes: (1) the female left the nest, (2) the male entered the box with food, and the female then used specific call types, (3) mates stopped calling but did not leave or enter the nest. The structure of vocal exchanges was globally stable between contexts, but females used calls with an up‐shifted spectrum during exchanges, at the end of which they left the nest or the male entered the nest. Birds vocalized more and at higher tempo during exchanges that ended up in feeding inside the nest. Birds also vocalized more during exchanges taking place during laying – a period of active mate guarding – than during incubation. We conclude that vocal exchanges could signal the females’ need for food and the males’ mate guarding behaviour, and discuss other possible functions of this communication.     

Sex roles, parental effort and offspring desertion in the monogamous Eurasian Curlew Numenius arquata

The reasons for female desertion of offspring and the evolution of predominantly male care among monogamous bird species are not clearly understood. We studied parental effort during the incubation and chick rearing periods in the Eurasian Curlew Numenius arquata in western Finland, and compared timing of brood desertion with other populations in Europe. Males and females contributed equally to incubation and showed no differences in the intensity of mobbing behaviour towards a potential nest predator (stuffed crow) shortly after hatching. However, females deserted their offspring approximately halfway through the brooding period ( c. 16 d after hatching), while males remained with chicks until independence ( c. 35 d). Females with late-laid clutches deserted their offspring sooner after hatching than those with clutches produced earlier in the season. Curlew females deserted younger chicks in northeast Europe, where laying dates were later, breeding seasons shorter and migration distances were longer, than in western and central Europe. We suggest that the most likely reasons for offspring desertion by females may be associated with increased female survivorship and maintenance of pairbond between years.     

Increasing the perceived predation risk changes parental care in female but not in male Great Tits Parus major

In birds, little is known about how the presence of predators alters parental food distribution decisions among nestlings. We found that experimentally increasing perceived predation risk changed parental care in female but not in male Great Tits Parus major. Females fed the lightest and average nestlings at similar rates under control conditions when predation risk was not manipulated but ignored the lightest nestling under increased perceived predation risk. Moreover, females reduced the duration of nest visits greatly after encountering a model predator, suggesting that the perception of predators may facilitate brood reduction mechanisms.     

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.     

A new method for catching cavity‐nesting birds during egg laying and incubation

ABSTRACT The physiological condition of female birds during the egg‐laying and incubation periods is of considerable interest and yet is relatively understudied in wild birds, primarily due to the difficulty of catching birds during this period without causing nest desertion. We therefore developed a box‐net to capture cavity‐nesting birds using sections of a mist‐net placed around a metal cubic frame. We captured female Great Tits (Parus major) as they left nest boxes during the egg‐laying and incubation periods and measured desertion rates. Using box‐nets, we captured 108 of 119 (90%) females during egg laying and 10 of 12 (83%) during incubation. Our recapture rate over two consecutive days during incubation was 50% (5 of 10). Females not captured left nest boxes before we attempted to capture them, escaped through a hole in the mist‐net, or remained in nest boxes for more than 2 h, after which we ended capture attempts. Overall, 22% of egg‐laying females deserted, with desertion rates highest early in the egg‐laying period. Desertion rates of females captured using box‐nets did not differ from those of undisturbed females. One of 10 females captured in a box‐net deserted during the incubation period. Box‐nets are portable, can be set up and taken down quickly and easily, and could potentially be used with nest boxes or natural cavities at any height. Box‐nets are easy to construct and adaptable for use with an array of cavity‐nesting birds, and can be an important tool for studying female physiology during egg laying and incubation.     

Female great tits Parus major do not increase their daily energy expenditure when incubating enlarged clutches

Several studies have shown that enlargement of clutches during incubation reduces the long-term survival of parents. In line with these findings, studies on the energetic costs of nocturnal incubation show an increase in energy expenditure with clutch enlargement. Studies on daily energy expenditure during incubation (DEE_inc), however, do not consistently show such a negative effect of clutch enlargement. To determine whether differential survival results from a direct increase in energy costs or rather from costs associated with compensatory behaviour, we studied the DEE_inc (kJ day⁻¹), change in body mass and nest attendance behaviour of free-living female great tits Parus major that incubated either control or experimentally enlarged clutches. DEE_inc did not differ between the two treatment groups, but was negatively related to mean ambient temperature over the 24-h measuring period, and to the fraction of daytime females spent on the nest. Controlling for these two factors, females incubating enlarged clutches did not spend more energy per 24 h period than females incubating control clutches. Clutch enlargement also did not affect body mass of incubating females, or their nest attendance behaviour. Yet, in the enlarged group body mass change and nest attentiveness were negatively correlated, suggesting that females responded differently to the experimental treatment and thereby preventing us from finding an effect of clutch enlargement.     

Costs of large communal clutches for male and female Greater Rheas Rhea americana

The breeding system of the Greater Rhea Rhea americana is almost unique among birds as it combines harem polygyny and sequential polyandry, with communal egg-laying and uniparental male care. In this species, large communal clutches (more than 30 eggs) are rare and have a lower hatching success than smaller clutches. Here we analyse the proximate causes of hatching failures and the costs of large communal clutches (and therefore the costs of extensive polygyny) for males and females. We evaluated if length of the nesting period, egg viability, egg losses during incubation and male parental activity at the nest were affected by clutch size. We also evaluated if chicks hatched from large clutches have a lower survival during the first 2 months after hatching. Large clutches had longer nesting period and lower hatching success, mainly as a result of bacterial contamination of the eggs and increased hatching asynchrony. In addition, large clutches tended to lose more eggs as a result of accidental breakage or predation. Male activity at the nest and chick survival were not related to clutch size. Low hatching success, nest predation risk and energetic costs associated with large clutches penalize females that join large harems and males that accept additional eggs into the nest.     

Incubation and brooding rhythm of the Cape Petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica

Incubation and brooding performance of the Cape Petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica, in the 1991 -1992 austral summer is reported in detail and compared with data from two other areas. There was an inverse relationship between the mean foraging trip length and mean peak weight of chicks. Variation in shift length throughout incubation resulted from a combination of seasonal effects (food supply) and behavioural adjustment to hatching. The length of the last incubation shift was independent of its number but decreased with date of initiation. The average weight of nestattending birds increased steadily during the incubation period and at the same rate in both sexes. Females attained seasonal peak body-weight before the egg hatched, whereas males just recovered their pre-breeding weight. Males took on a slightly larger share of incubation (52%, range 40–63%) and invested more time in mutual nest attendance. Length of foraging trips varied consistently in pairs and individuals, its repeatability being twice as high in females as in males. The incubation performance of individual females (length of the first and an average foraging trip) was correlated with the size of the egg they had laid and also to subsequent chick growth and fledging success. There was a tendency in successful pairs to spend more time together at the nest and to have shorter foraging trips. I suggest that variation in breeding performance among pairs was mainly a result of individual variation in female quality, with pairs where the female contributed more to incubation being more successful. There is little evidence that egg losses were caused by parental errors.     

A single long day triggers follicle growth in captive female great tits (Parus major) in winter but does not affect laying dates in the wild in spring

In many forest passerine bird species, rapid climate warming has led to a phenological mismatch between the period of maximum nestlings' food requirements and the period of maximum food availability (seasonal caterpillar biomass peak) due to an insufficient advancement of the birds' laying dates. The initiation of laying is preceded by the development of the gonads, which in birds are regressed outside the breeding season. Increasing day length in late winter and early spring triggers a cascade of hormones which induces gonadal development. Since day length is not altered by climate change, one potential restriction to advancing laying date is the seasonal timing of gonadal development. To assess the importance of gonadal growth for timing of reproduction we experimentally manipulated the timing of gonadal development. We show that the growth of the largest follicle of captive female great tits (Parus major) increased after being exposed to just a single long day in winter (20 hours of light followed by 4 hours darkness). We then photostimulated wild female great tits from two study areas in a field experiment in spring for a single day and determined their laying date. These populations differed in the availability of food allowing us to test if food availability in combination with photostimulation affected egg laying dates. Despite an expected difference in the onset of gonadal growth, laying dates of photostimulated females did not differ from control females in both populations. These results suggest that wild great tits are not restricted in the advancement of their laying date by limited gonadal development.     

Why do Both Parents Incubate in the Kentish Plover?

Incubation by both parents is a common parental behaviour in many avian species. Biparental incubation is expected if the survival prospects of offspring are greatly raised by shared care, relative to the costs incurred by each parent. We investigated this proposition in the Kentish plover Charadrius alexandrinus, in which both parents incubate the clutch, but one parent (either the male or the female) usually deserts after hatching of the eggs. We carried out a mate‐removal and food supplementation experiment to reveal both the role of the sexes and food abundance in maintaining biparental incubation by removing either the male or the female from the nest for a short period of time. In some nests we provided supplementary food for the parent that remained at the nest to reduce the costs of incubation, whereas other nests were left unsupplemented. Although males spent more time on incubation after their mate had been removed, females’ incubation did not change. Notwithstanding the increased male incubation, total nest attentiveness was lower at uniparental nests than at biparental controls. However, incubation behaviour was not influenced by food supplementation. We conclude that offspring desertion during incubation is apparently costly in the Kentish plover, and this cost cannot be ameliorated with supplementary food.     

Predator-Specific Effects on Incubation Behaviour and Offspring Growth in Great Tits

In birds, different types of predators may target adults or offspring differentially and at different times of the reproductive cycle. Hence they may also differentially influence incubation behaviour and thus embryonic development and offspring phenotype. This is poorly understood, and we therefore performed a study to assess the effects of the presence of either a nest predator or a predator targeting adults and offspring after fledging on female incubation behaviour in great tits (Parus major), and the subsequent effects on offspring morphological traits. We manipulated perceived predation risk during incubation using taxidermic models of two predators: the short-tailed weasel posing a risk to incubating females and nestlings, and the sparrowhawk posing a risk to adults and offspring after fledging. To disentangle treatment effects induced during incubation from potential carry-over effects of parental behaviour after hatching, we cross-fostered whole broods from manipulated nests with broods from unmanipulated nests. Both predator treatments lead to a reduced on- and off-bout frequency, to a slower decline in on-bout temperature as incubation advanced and showed a negative effect on nestling body mass gain. At the current state of knowledge on predator-induced variation in incubation patterns alternative hypotheses are feasible, and the findings of this study will be useful for guiding future research.     

Effects of a food supplementation experiment on reproductive investment and a post-mating sexually selected trait in magpies Pica pica

Food availability is an important factor affecting breeding success in birds. Food supplementation experiments in birds have in general focused on the effects on reproductive success in terms of female investment (laying date, clutch size, egg size), however, it is also known that the estimation of mate quality based on sexually selected signals influences female reproductive investment. In the particular case of magpies, females use nest size, a post-mating sexually selected signal, to assess male's likelihood to invest in reproduction, and accordingly adjust reproductive investment (clutch size). Then, the possible effects of food supplementation on female reproductive investment could be mediated by other variables related to parental quality, such as nest size in magpies. In the present study, we explore if higher food availability in a magpie territory affected both male sexually selected traits (i.e. nest size) and female reproductive investment (laying date, egg size, clutch size). We performed a food supplementation experiment in which we experimentally increased food availability in several magpie territories, keeping others as controls. In food-supplemented territories, males built significantly larger nests and females significantly increased egg size by 4.1% compared to control females. Results suggest that the continuous provisioning of protein rich food allowed magpie females to increase egg size. However, laying date and clutch size did not differ between control and food-supplemented magpie pairs. Food availability also affected the relationship between female reproductive investment and nest size. In control territories, females decreased their egg size in response to a larger nest, whereas a tendency for the opposite relationship was revealed in food-supplemented territories. We discuss the possibility that magpie females adopt different strategies for reproductive investment according to food availability.     

Does food supplementation really enhance productivity of breeding birds?

Food availability influences multiple stages of the breeding cycle of birds, and supplementary feeding has helped in its understanding. Most supplementation studies have reported advancements of laying, whilst others, albeit less numerous, have also demonstrated fitness benefits such as larger clutches, shorter incubation periods, and greater hatching success. Relatively few studies, however, have investigated the effects of supplementary feeding for protracted periods across multiple stages of the breeding cycle. These effects are important to understand since long-term food supplementation of birds is recommended in urban habitats and is used as a tool to increase reproductive output in endangered species. Here, we compare the breeding phenology and productivity of blue tits Cyanistes caeruleus and great tits Parus major breeding in food-supplemented and non-supplemented blocks in a broadleaf woodland in central England over three seasons (2006–2008). Supplementation was provided continuously from several weeks pre-laying until hatching, and had multiple significant effects. Most notably, supplementation reduced brood size significantly in both species, by half a chick or more at hatching (after controlling for year and hatching date). Reduced brood sizes in supplemented pairs were driven by significantly smaller clutches in both species and, in blue tits, significantly lower hatching success. These are novel and concerning findings of food supplementation. As expected, supplementary feeding advanced laying and shortened incubation periods significantly in both species. We discuss the striking parallels between our findings and patterns in blue and great tit reproduction in urban habitats, and conclude that supplementary feeding may not always enhance the breeding productivity of birds.     

Handicapped females receive more feedings during incubation from their mates: support for the female nutrition hypothesis

The female nutrition hypothesis posits that provisioning intensity of incubating females by their mates may depend on female needs and ensure proper incubation and a corresponding high hatching and breeding success of breeding pairs. Here, we have handicapped female pied flycatchers Ficedula hypoleuca at the beginning of incubation by clipping two primaries on each wing and filmed nests during incubation and later nestling provisioning to estimate male involvement in incubation feeding at the nest and in offspring care. Incubation feeding was more frequent at late nests. Correcting for this seasonal effect, incubation feeding was significantly affected by treatment and twice as high at experimental as at control nests. There was no effect of the experiment on female incubation attendance. The handicap did not result in any effect on hatching and breeding success, nestling growth and male or female provisioning and mass at the end of the nestling period. Males adjust their incubation feeding activity at the nest to female energetic requirements during incubation.     

On the origin and rarity of interspecific nest parasitism in birds

Interspecific nest parasitism is surprisingly rare in birds given the potential advantages for the parasite of exploiting the parental care of other species. One possibility is that chicks will not thrive with the parental care and food of heterospecifics. I simulated parasitism in nonparasitic congeners by switching eggs between nests of three species of titmice (great tit Parus major, blue tit Parus caeruleus, and coal tit Parus ater). The experiment showed that compatibility of parental care was not a constraint preventing parasitism. I also used the model system to compare fitness consequences of inter- and intraspecific nest parasitism, addressing the problem of which form is ancestral. Fledging success (body mass, survival) was higher when an egg was added to the nest of a smaller species than to the nest of a conspecific and also higher when the parasitic chick hatched early rather than late relative to host chicks. This suggests that interspecific nest parasitism may not require a stage of intraspecific nest parasitism before evolving but may start from a larger species directly exploiting the parental care of a smaller species or a species with shorter incubation period directly exploiting a species with longer incubation period.     

Egg speckling patterns do not advertise offspring quality or influence male provisioning in great tits

Many passerine birds lay white eggs with reddish brown speckles produced by protoporphyrin pigment. However, the function of these spots is contested. Recently, the sexually selected eggshell coloration (SSEC) hypothesis proposed that eggshell color is a sexually selected signal through which a female advertises her quality (and hence the potential quality of her future young) to her male partner, thereby encouraging him to contribute more to breeding attempts. We performed a test of the SSEC hypothesis in a common passerine, the great tit Parus major. We used a double cross-fostering design to determine whether males change their provisioning behavior based on eggshell patterns they observe at the nest. We also tested the assumption that egg patterning reflects female and/or offspring quality. Because birds differ from humans in their color and pattern perception, we used digital photography and models of bird vision to quantify egg patterns objectively. Neither male provisioning nor chick growth was related to the pattern of eggs males observed during incubation. Although heavy females laid paler, less speckled eggs, these eggs did not produce chicks that grew faster. Therefore, we conclude that the SSEC hypothesis is an unlikely explanation for the evolution of egg speckling in great tits.