The cost of incubation in relation to clutch-size in the Collared Flycatcher Ficedula albicollis

This study examines the importance of avian incubation costs as determinants of clutch-size variation by performing clutch-size and brood-size manipulations in the same population of Collared Flycatchers Ficedula albicollis during the same breeding season. In 2 5 cases when three or more clutches of the same size were completed on the same day, we moved two eggs on the day after the last egg had been laid from one randomly selected clutch (C) to another (C) and moved two other eggs from this to a third clutch (C⁺). In 20 other cases of simultaneously completed clutches of the same size, we moved two randomly selected young from one brood to a second and from that moved two other young to a third (B, B and B⁺groups). Most females were weighed the day after completion of the clutch and 1–4 days before hatching of the young, and some of them also 10–14 days after hatching of the young. We measured the daily energy expenditure of females incubating manipulated clutches of 4, 6 and 8 eggs by means of the doubly-labelled water (D₂¹⁸O) technique and also recorded their nest attendance. Hatching success of fertilized eggs was reduced in the enlarged clutches compared with control and reduced clutches. Females expired on average 3142.6 ml CO₂ and expended 78.6 kJ per day while incubating, which corresponds to a metabolic intensity of 3.3 times BMR. Daily energy expenditure increased with clutch-size due to higher costs while incubating, and not because of changed activity patterns. There were no significant differences in length of incubation, female mass or mass changes between phases for the C, C and C⁺groups. In both the C and B groups, enlarged broods produced significantly more fledged young than control broods, and those significantly more than reduced broods. Fledgling tarsus-length and mass did not differ significantly between treatments in either the C or B groups. There was no significant difference in breeding success between clutch and brood manipulations. In this season, incubation costs did not entail significant fitness losses, expressed either as fledgling production or female condition. Also, control females could have raised more young to fledging age than they did with no apparent costs.     

Clutch size manipulation, hatching success and offspring phenotype in the ball python (Python regius)

In a diverse array of avian and mammalian species, experimental manipulations of clutch size have tested the hypothesis that natural selection should adjust numbers of neonates produced so as to maximize the number of viable offspring at the end of the period of parental care. Reptiles have not been studied in this respect, probably because they rarely display parental care. However, females of all python species brood their eggs until hatching, but they do not care for their neonates. This feature provides a straightforward way to experimentally increase or reduce clutch size to see whether the mean clutch size observed in nature does indeed maximize hatching success and/or optimize offspring phenotypes. Eggs were removed or added to newly laid clutches of Ball Pythons ( Python regius ) in tropical Africa (nine control clutches, eight with 50% more eggs added, six with 42% of eggs removed). All clutches were brooded by females throughout the 2-month incubation period. Experimental manipulation of clutch-size did not significantly affect the phenotypes (morphology, locomotor ability) of hatchlings, but eggs in 'enlarged' clutches hatched later, and embryos were more likely to die before hatching. This mortality was due to desiccation of the eggs, with females being unable to cover 'enlarged' clutches sufficiently to retard water loss. Our results support the notion of an optimal clutch size, driven by limitations on parental ability to care for the offspring. However, the proximate mechanisms that generate this optimum value differ from those previously described in other kinds of animals. © 2003 The Linnean Society of London . Biological Journal of the Linnean Society 2003, 78 , 263–272.     

Does the cost of incubation set limits to clutch size in common eiders Somateria mollissima?

We examined the effect of natural clutch size on the cost of incubation in a population of common eiders Somateria mollissima nesting in Tromsø, northern Norway. The body condition of females at day 5 in the incubation period was not related to clutch size (3–6 eggs), but females incubating large clutches lost more mass and had a lower body condition at day 20 in the incubation period than females incubating small clutches. Females incubating large clutches had a slightly shorter incubation period and a lower egg predation rate. The results do not support the hypothesis that the female's ability to produce eggs is the only ultimate control of clutch size in eider. Instead the results suggest that there may be an interaction between the allocation of body reserves to eggs and incubation, and that females producing large clutches allocate more of their body reserves to incubation than females producing small clutches, in order to shorten the incubation period and to minimise the risk of predation on eggs.     

Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

Lack ( 1967 ) proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per‐capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two‐egg clutches to 0.50 for seven‐egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three‐egg clutches to 27.7 days (95% CI 27.3–28.1) for six‐egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five‐egg clutch.     

Hatching asynchrony that maintains egg viability also reduces brood reduction in a subtropical bird

In birds, hatching failure is pervasive and incurs an energetic and reproductive cost to breeding individuals. The egg viability hypothesis posits that exposure to warm temperatures prior to incubation decreases viability of early laid eggs and predicts that females in warm environments minimize hatching failure by beginning incubation earlier in the laying period, laying smaller clutches, or both. However, beginning incubation prior to clutch completion may incur a cost by increasing hatching asynchrony and possibly brood reduction. We examined whether Florida scrub jays (Aphelocoma coerulescens) began incubation earlier relative to clutch completion when laying larger clutches or when ambient temperatures increased, and whether variation in incubation onset influenced subsequent patterns of hatching asynchrony and brood reduction. We compared these patterns between a suburban and wildland site because site-specific differences in hatching failure match a priori predictions of the egg viability hypothesis. Females at both sites began incubation earlier relative to clutch completion when laying larger clutches and as ambient temperatures increased. Incubation onset was correlated with patterns of hatching asynchrony at both sites; however, brood reduction increased only in the suburbs, where nestling food is limiting, and only during the late nestling period. Hatching asynchrony may be an unintended consequence of beginning incubation early to minimize hatching failure of early laid eggs. Food limitation in the suburbs appears to result in increased brood reduction in large clutches that hatch asynchronously. Therefore, site-specific rates of brood reduction may be a consequence of asynchronous hatching patterns that result from parental effort to minimize hatching failure in first-laid eggs. This illustrates how anthropogenic change, such as urbanization, can lead to loss of fitness when animals use behavioral strategies intended to maximize fitness in natural landscapes.     

Phytohemagglutinin response and immunoglobulin index decrease during incubation fasting in female common eiders

To maximize their fitness, long-lived species face trade-offs between survival and reproduction. The cost of reproduction, which is defined as the negative effect of current parental investment on chances of adult survival and future reproduction, may affect immune function, possibly through hormonal changes. In this study, components of acquired immunity and plasma corticosterone levels of female eiders (Somateria mollissima) have been measured throughout the incubation period as a function of clutch size. These precocial birds lay up to six eggs and fast completely during incubation. Birds were sampled early and late in the incubation period, with clutches ranging from one to four eggs. T-cell-mediated immune response and humoral immunity were assessed by phytohemagglutinin (PHA) skin tests (a challenging method) and measurements of serum immunoglobulins (a monitoring method), respectively. During incubation, responses to PHA injection and immunoglobulin index significantly decreased, by about 40% and 25%, respectively. These observed decreases occurred independently of the number of eggs laid by the females. Corticosterone did not vary significantly during incubation, whatever the clutch size. We conclude that female eiders seem to reallocate their resources from immune function to reproductive effort independently of clutch size or corticosterone levels.     

Females Paired with New and Heavy Mates Reduce Intra-Clutch Differences in Resource Allocation

Reproductive investment affects both offspring and parental fitness and influences the evolution of life histories. Females may vary their overall primary reproductive effort in relation to the phenotypic characteristics of their mate. However, the effects of male quality on differential resource allocation within clutches have been largely neglected despite the potential implications for mate choice and population dynamics, especially in species exhibiting biparental care and brood reduction. Female southern rockhopper penguins Eudyptes chrysocome paired with heavy mates reduced intra-clutch variation in egg and albumen masses. Females paired with new mates also reduced intra-clutch variation in yolk androgen levels. Since both an increased mass and increased androgen concentrations positively influence chick survival under sibling competition, the chances of fledging the whole clutch are likely to be higher for newly formed pairs with heavy males than for previously formed pairs with light males. Interestingly, total clutch provisioning did not vary with male quality. We show for the first time that females vary intra-clutch variation in resource allocation according to male quality. In species with brood reduction, it may be more adaptive for females to modulate the distribution of resources within the clutch according to breeding conditions, than to change their total clutch provisioning.     

Maternal behavior and clutch manipulation in the ring-legged earwig (Dermaptera: Carcinophoridae)

The expression and maintenance of maternal behavior in the earwig,Euborellia annulipes, was examined through manipulation of clutch size, age, and species and through observations of interactions between brooding females. Females underwent discrete gonadotrophic cycles culminating in oviposition of first clutches that were highly variable in size. Neither the head capsule width nor the age of the mother was correlated with clutch size. Maternal care extended through embryogenesis and for the week following hatching. Clutch removal significantly shortened the interclutch interval, indicating that the presence of brood inhibited the onset of the second gonadotrophic cycle. Brooding females readily accepted replacement clutches of the same age. Thus, mothers did not appear to distinguish their own eggs from those of other females. Experimental doubling of clutch size did not significantly reduce the proportion hatching or fledging. In contrast, reducing clutch size diminished the percentage successfully fledging. Manipulation of clutch age resulted in reduced hatching/fledging success. Placing two females, each with newly laid clutches, in the same cage usually resulted in egg transfer from the nest of one female to that of the other within 12 h. Nests of females with larger forceps were significantly more likely to contain both clutches. When mothers with first clutches were paired with mothers with third clutches, eggs were more likely to be transferred to the nest of the older female.E. annulipes females with newly laid clutches appeared to accept as replacement clutches eggs of the earwigDoru taeniatum. Alien clutches were maintained for the typical duration of embryogenesis; however, noD. taeniatum hatchlings were observed.     

Determination of clutch-size in the Kentish Plover Charadrius alexandrinus

We studied the clutch-size distribution of Kentish Plover Charadrius alexandrinus in 1991 and 1992 and tested three hypotheses for determination of clutch-size: egg-formation ability of females, incubation ability of parents and nest predation. Variation in clutch-size was small: 71 out of 74 clutches had three eggs (coeff. var. = 7%). Females spent more time foraging (51 ± 6%) pre-laying and during egg-laying than their mates (39 ± 5%). However, we concluded that egg-formation was not constrained by food availability because 77% and 100% of clutches were initiated before the peaks of prey density and of prey mass, respectively. Furthermore, the number of clutches initiated and completed over time was unrelated to prey density. By experimentally reducing and enlarging clutches, we found that enlarged clutches of four eggs took longer to hatch (24.8 ± 0.9 days) than control clutches of three (21.6 ± 0.7 days). Eggs of enlarged clutches also lost weight more slowly during incubation in both years compared with control clutches. No difference was found in the incubation behaviour or weight loss of parents between reduced, control and enlarged clutches. We found no evidence to support the nest predation hypothesis, since neither the proportion of nests predated nor the number of chicks hatched was different between reduced, control and enlarged clutches. The results of this study are most consistent with the incubation ability hypothesis, although the parental ability hypothesis has remained untested.     

Paternal care and male mate-attraction effort in the European starling is adjusted to clutch size

In facultative polygynous birds with biparental care, a trade-off may occur between male parental care and attraction of additional mates. If there is a cost associated with reduced male parental care, the relative benefit of mate attraction may be predicted to decrease as the size of a male's clutch or brood increases. We tested this prediction in monogamous pairs of facultatively polygynous European starlings (Sturnus vulgaris). The larger the clutch, the more time the male spent incubating and the less time he spent attracting an additional female (i.e. singing near and carrying green nesting material into adjacent empty nest-boxes). Reduced paternal incubation resulted in lower overall incubation (the female did not compensate) and lower hatching success. Immediately after experimental reduction of clutches, males spent significantly less time incubating and more time singing and carrying greenery, and vice versa for experimentally enlarged clutches. Males with experimentally reduced clutches attracted a second female more often than males with experimentally enlarged clutches. This is the first study, to our knowledge, to provide experimental evidence for an adjustment of paternal care and male mate-attraction effort to clutch size. However, a trade-off between paternal nestling provisioning and mate attraction was not revealed, probably due to the absence of unpaired females by that time in the breeding season. Experiments showed that the relative contribution of the male and female to nestling provisioning was unrelated to brood size.     

Effect of clutch size on male egg-fanning behavior and hatching success in the goby, Eviota prasina (Klunzinger)

In fish that exhibit paternal care, the females often choose their mates on the basis of male traits that are indicative of the parental ability of the males. In a marine goby, Eviota prasina, males tend their eggs within their nests until hatching, and females prefer males that have longer dorsal fins and exhibit courtship behavior with a higher frequency as their mates. In order to clarify the relationship between these sexually selected traits and the parental ability of males of E. prasina, the factors affecting the hatching success of eggs within male nests and the male parental care behavior were examined in an aquarium experiment. Females spawned their eggs in male nests and the clutch size of females showed a high individual variation (range = 88-833 eggs). The hatching success of eggs within male nests showed a positive correlation with the time spent by males in fanning eggs and the clutch size. In contrast to the prediction, however, the hatching success did not show a significant correlation with the sexually selected traits, i.e., the male dorsal fin length and the frequency of courtship displays. Moreover, multiple regression analysis indicated that the time spent by the males in fanning was the most important factor affecting the survival rate of the eggs. The time spent by males in fanning behavior was influenced by the clutch size within their nests; the fanning behavior of males occurred with a higher frequency when they tended larger clutches. Males are required to invest a greater effort in egg-tending behavior to achieve a higher hatching success when they receive larger clutches, probably due to the greater reward for their parental behavior. Based on their mate choice, females may obtain other benefits such as high quality offspring.     

Time-dependent reproductive decisions in the blue tit

Many breeding attempts in birds do not result in any fledged young due to predation on eggs or young. Consequently, the influence of time constraints on reproductive decisions are integrated parts of the reproductive behaviour of birds breeding within short, seasonal climate zones. In this study, I mimicked nest predation by removing blue tit (Parus caeruleus) clutches shortly after completion. Around 75% of the removed clutches were followed by a repeat clutch. Females producing their first clutch early in the season and females with an early onset of incubation in the laying sequence (an indication of high parental or territory quality) were most likely to initiate a repeat clutch. A trade‐off between the benefits of a repeat clutch and survival likely stopped late females in bad condition from investing more in the current reproductive season. Females producing a repeat clutch laid fewer eggs, had an earlier onset of incubation in the laying sequence and produced larger eggs than they did when producing their original first clutch. Eggs produced after the onset of incubation were especially large in the repeat clutches. Since food availability was presumably higher when the female produced her repeat clutch compared with her first clutch, females made a strategical decision when reducing clutch size, whereas onset of incubation and egg size may have been energetically constrained when producing the first clutch. Females that produced a relatively large clutch, had a relatively early onset of incubation, and laid relatively large eggs in their first clutch also did so when producing a repeat clutch, indicating that some of the variation in breeding parameters are due to differences in parental or territory quality. Differences between years in the temperature‐dependent development rate of caterpillars seem to affect the time constraints on breeding. A year with a predicted early seasonal decline in caterpillars resulted in short intervals between removal and relaying, small clutches and an early onset of incubation.     

Cost of reproduction in a long-lived bird: incubation effort reduces immune function and future reproduction

Life-history theory predicts that increased current reproductive effort should lead to a fitness cost. This cost of reproduction may be observed as reduced survival or future reproduction, and may be caused by temporal suppression of immune function in stressed or hard-working individuals. In birds, consideration of the costs of incubating eggs has largely been neglected in favour of the costs of brood rearing. We manipulated incubation demand in two breeding seasons (2000 and 2001) in female common eiders (Somateria mollissima) by creating clutches of three and six eggs (natural range 3-6 eggs). The common eider is a long-lived sea-duck where females do not eat during the incubation period. Mass loss increased and immune function (lymphocyte levels and specific antibody response to the non-pathogenic antigens diphtheria and tetanus toxoid) was reduced in females incubating large clutches. The increased incubation effort among females assigned to large incubation demand did not lead to adverse effects on current reproduction or return rate in the next breeding season. However, large incubation demand resulted in long-term fitness costs through reduced fecundity the year after manipulation. Our data show that in eiders, a long-lived species, the cost of high incubation demand is paid in the currency of reduced future fecundity, possibly mediated by reduced immune function.     

Blue–Green Eggs in Pied Flycatchers: An Experimental Demonstration that a Supernormal Stimulus Elicits Improved Nestling Condition

It has been proposed that blue–green egg colouration in certain avian species has evolved as a signal of female and egg quality to males, affecting their investment in the brood according to the differential allocation scenario. A recent experiment has successfully manipulated male investment by placing dummy eggs simulating the extremes in the blue–green natural reflectance. Here we have substituted one egg in certain pied flycatcher clutches with a single deep‐blue (blue) or light‐blue (pale) dummy egg and used other clutches as controls. Blue dummy eggs reflected more in the blue–green part of the spectrum than natural and pale dummy eggs, which were similar in reflectance. Nestlings in nests with one blue dummy egg during laying and incubation attained a significantly higher mass and condition than those in nests with one pale dummy egg or in control broods, when controlling for phenology, brood size and blue–green chroma of the clutch. Only a shift in parental investment can have induced this highly significant effect. Preference for deeply coloured clutches associated with high‐quality females and their maternal effects may explain stimulation by a single supernormal stimulus emanating from multiple objects like clutches.     

Cascading costs of reproduction in female house wrens induced to lay larger clutches

In many species, females produce fewer offspring than they are capable of rearing, possibly because increases in current reproductive effort come at the expense of a female's own survival and future reproduction. To test this, we induced female house wrens (Troglodytes aedon) to lay more eggs than they normally would and assessed the potential costs of increasing cumulative investment in the three main components of the avian breeding cycle – egg laying, incubation and nestling provisioning. Females with increased clutch sizes reared more offspring in the first brood than controls, but fledged a lower proportion of nestlings. Moreover, nestlings of experimental females were lighter than those of control females as brood size and prefledging mass were negatively correlated. In second broods of the season, when females were not manipulated, experimental females laid the same number of eggs as controls, but experienced an intraseasonal cost through reduced hatchling survival and a lower number of young fledged. Offspring of control and experimental females were equally likely to recruit to the breeding population, although control females produced more recruits per egg laid. The reproductive success of recruits from broods of experimental and control females did not differ. The manipulation also induced interseasonal costs to future reproduction, as experimental females had lower fecundity than controls when breeding at least 2 years after having their reproductive effort experimentally increased. Finally, females producing the modal clutch size of seven eggs in their first broods had the highest lifetime number of fledglings.     

More colourful eggs induce a higher relative paternal investment in the pied flycatcher Ficedula hypoleuca: a cross-fostering experiment

The signalling hypothesis of eggshell colouration in birds posits that females of species with blue-green eggs signal their phenotypic quality or the quality of their eggs to their mates through deposition of the antioxidant biliverdin as pigment. Males respond by investing more in the offspring. Through a cross-fostering experiment where we have exchanged whole clutches between pairs of pied flycatcher Ficedula hypoleuca nests, we managed to break potential associations between female quality and clutch chromaticity. We show that males respond to incubated clutches with more variable and higher peak values in blue-green chroma through a higher proportional investment in nestling provisioning on day 4 of the nestling period, when males invest more heavily than females in provisioning. More variable clutches show higher peak chroma values. We also show that egg colour during the two-week incubation period has a significant effect, which is not found for the colour of eggs during the laying period. Finally, the proportion of male provisioning visits affects negatively female brooding effort and nestling mortality, and thereby has positive effects on female fitness. Blue-green chroma in the pied flycatcher functions as a signal of female or clutch quality to males which respond by adjusting their relative investment with respect to total pair effort.     

Egg-size variation in the bluethroat (Luscinia s. svecica): constraints and adaptation

We examined inter- and intra-clutch egg-size variation in the bluethroat (Luscinia s. svecica), an open-nesting passerine breeding in the sub-alpine region in southern Norway. By removing first clutches shortly after egg-laying, we induced laying of a repeat clutch. Females significantly reduced the number of eggs from the first to the second nesting attempt, but increased mean egg size. Females in good condition laid significantly larger eggs than those in poor condition. Consistent with predictions of the brood survival hypothesis, assuming an adaptive investment in last eggs to ensure survival of all eggs in the clutch, we found that the size of the last eggs in first clutches was generally larger than the mean egg size of the clutch, and that the relative size of the last egg increased with clutch size. However, a large last egg reflected a general increase in egg size throughout the laying sequence rather than a specific investment in the last egg only. Egg size was not significantly influenced by sex or paternity (within-pair versus extra-pair) of the embryo. In repeat clutches the last egg was not consistently larger than the mean for the clutch. We conclude that female bluethroats face resource limitations during egg formation early in the season, and that the patterns of increase in egg size with laying order for first clutches, and from first to repeat clutches, can largely be explained by proximate constraints on egg formation.     

CLUTCH-SIZE, INCUBATION AND HATCHING SUCCESS IN THE HOUSE SPARROW AND TREE SPARROW PASSER SPP. AT OXFORD

Clutch-size, incubation and hatching success were studied in P. domesticus and P. montanus in 1961 and 1963–64 at Oxford. The most frequent clutch-size was four eggs in P. domesticus and five eggs in P. montanus . With one exception, colonies of P. domesticus showed no significant annual or local variations in its mean clutch-size; in P. montanus , however, there were significant annual variations in the mean clutch-size. Both species showed a seasonal increase followed by a decrease in their mean clutch-sizes.
Partial incubation occurred during the laying period of the clutch; sufficient incubation for continuous development of the embryo was apparently achieved when the last egg had been laid in clutches of two and three eggs in P. domesticus and in clutches of four eggs in P. montanus , but when the penultimate egg had been laid in larger clutches of both species. On average, hatching in P. domesticus occurred more or less synchronously in all eggs in clutches of two and three eggs, and in all eggs except the last one laid in larger clutches; the last egg in the larger clutches hatched up to a day after the others. It is suggested that this pattern of hatching was brought about by the pattern of incubation during the laying period.
P. domesticus had a lower hatching success than P. montanus , probably because fewer of its eggs were fertile.     

Reproductive effort reduces long-term immune function in breeding tree swallows (Tachycineta bicolor)

We examined whether strategies of reproductive allocation may reduce long-term immunocompetence through the effects of manipulated effort on secondary or acquired immunity. We tested whether increased reproductive effort leads to reduced immune function and survival by manipulating brood size in tree swallows (Tachycineta bicolor) and exposing breeding females to a primary and secondary exposure of sheep red blood cells to elicit a humoral immune response. Females raising enlarged broods produced fewer secondary antibodies than did females raising control or reduced broods. Most importantly, individuals with high secondary responses were more likely to survive to breed 3 years after brood manipulations, suggesting that differences in disease susceptibility may be caused by trade-offs in reproductive allocation. We also found that individual quality, measured by clutch initiation date, mediated the effects of brood manipulations, with higher-quality birds showing a greater ability to deal with increases in effort.     

Mutual ornamentation and the parental behaviour of male and female Collared Flycatchers Ficedula albicollis during incubation

One of the benefits of mate choice based on sexually selected traits is the greater investment of more ornamented individuals in parental care. The choosy individual can also adjust its parental investment to the sexual signals of its partner. Incubation is an important stage of avian reproduction, but the relationship between behaviour during incubation and mutual ornamentation is unclear. Studying a population of Collared Flycatchers Ficedula albicollis, we monitored the behaviour of both sexes during incubation in relation to their own and their partner's plumage traits, including plumage‐level reflectance attributes and white patch sizes. There was a marginally positive relationship between male feeding rate during incubation and female incubation rate. Female but not male behavioural traits were associated with the laying date of the first egg and clutch size. The behaviour of the two sexes jointly determined the relative hatching speed of clutches and the hatching success of eggs. Females with larger white wing patches spent less time incubating eggs and left the nestbox more frequently. Males with larger white wing patches fed females less frequently, whereas males with brighter white plumage areas visited the nestbox more regularly without feeding. Females tended to leave the nest less often when mated to males with larger wing patches, and females spent less time incubating when males had more UV chromatic plumage. The behaviour of both partners during incubation therefore predicted hatching patterns and was correlated with their own and sometimes with their partner's plumage ornamentation. These results call for further studies of mutual ornamentation and reproductive effort during incubation.