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.     

Fitness consequences of male provisioning of incubating females in a desert passerine bird

Male provisioning of incubating females can increase reproductive success by maintaining physiological condition of females and consistency of incubation. The effects of male provisioning on the maintenance of incubation temperature and embryo development should be particularly pronounced in environments where ambient temperature exceeds the tolerance of unincubated eggs and where consistency of female incubation might be particularly important for hatching success. Here, we investigated the reproductive consequences of incubation feeding in a desert population of House Finches (Carpodacus mexicanus) in southwestern Arizona. We found that greater nest attentiveness by females was related to higher minimum incubation nest temperature, that in turn was closely associated with hatching success. Only 44% of males regularly provisioned their incubating females. Although provisioned females maintained higher incubation temperature and took fewer incubation breaks than non-provisioned females, overall, male provisioning did not influence incubation dynamics or hatching success. Further, a male’s incubation feeding rate did not correlate with male provisioning of nestlings. These results corroborate the finding that, in male House Finches, neither provisioning of incubating females nor pre-incubation courtship feeding are associated with increases in circulating pituitary prolactin––the hormone regulating male provisioning of nestlings. We suggest that incubation provisioning by male might be a component of pair maintenance behavior and that variation in male incubation behavior is best understood in relation to asymmetries in residual reproductive values between the mates.     

Incubation feeding by male Scarlet Tanagers: a mate removal experiment

ABSTRACT Incubation feeding, where males feed their mates, is a common behavior in birds and may improve female condition, nest attentiveness, and nesting success. We used behavioral observations and a temporary mate removal experiment to test the female nutrition hypothesis for incubation feeding by male Scarlet Tanagers (Piranga olivacea). All males (N= 20) were observed incubation feeding and fed females both at the nest (x? 1.36 trips/h) and away from the nest (x? 20.1 trips/h). Male feeding rate off‐nest was negatively correlated with the duration of female foraging bouts and positively correlated with the total time females spent incubating per hour. Eggs were predated at seven of 19 (37%) nests, but nest survival during incubation was not related to either female incubation behavior or male feeding rate. During temporary removal experiments (N= 12), female Scarlet Tanagers remained on the nest significantly longer and did not have longer foraging bouts. An unexpected outcome of the removal experiments was a dramatic change in female vocal behavior. All 12 experimental females gave chik‐burr calls during the male‐removal experiments (x? bout length = 11.7 min), but during normal observation periods only six of 20 females at the incubation stage gave chik‐burr calls (x? bout length = 0.7 min, N= 20). Our results suggest that female tanagers likely gain nutritional benefits from incubation feeding, but male feeding may not improve immediate reproductive success. Nine of 54 (17%) nestlings in five of 17 broods (29%) were extra‐pair young (EPY), indicating that males could potentially benefit from incubation feeding via mate retention and fidelity as well as, or instead of, through immediate gains in reproductive success. Our study indicates that females benefit from incubation feeding and do not simply passively accept food from their mates, but instead may influence male feeding rates through direct (e.g., mate following and vocalizing) and indirect (the threat of mate abandonment or cuckoldry) means.     

The Effect of Male Incubation Feeding,Food and Temperature on the Incubation Behaviour of New Zealand Robins

Because of finite resources, organisms face conflict between their own self‐care and reproduction. This conflict is especially apparent in avian species with female‐only incubation, where females face a trade‐off between time allocated to their own self‐maintenance and the thermal requirements of developing embryos. We recorded incubation behaviour of the New Zealand robin (Petroica longipes), a species with female‐only incubation, male incubation feeding and high nest predation rates. We examined how male incubation feeding, ambient temperature and food availability (invertebrate biomass) affected the different components of females’ incubation behaviour and whether incubation behaviour explained variation in nest survival. Our results suggest that male incubation feeding rates of 2.8 per hour affect the female’s incubation rhythm by reducing both on‐ and off‐bout duration, resulting in no effect on female nest attentiveness, thus no support for the female‐nutritional hypothesis. The incubation behaviours that we measured did not explain nest survival, despite high nest predation rates. Increased ambient temperature caused an increase in off‐bout duration, whereas increased food availability increased on‐bout duration. While males play a vital role in influencing incubation behaviour, female robins attempt to resolve the trade‐off between their own foraging needs and the thermal requirements of their developing embryos via alternating their incubation rhythm in relation to both food and temperature.     

Is Mate Provisioning Predicted by Ornamentation? A Test with Northern Cardinals (Cardinalis cardinalis)

Male birds of many species feed their mates during courtship and incubation. The amount of food provided can be substantial and even essential for successful reproduction in some species, and can influence female nest attentiveness in many others. Additionally, mate provisioning may predict later nestling feeding rates. Females may thus benefit from being able to determine male provisioning effort. We assessed the expression of several ornaments, known to indicate condition in male northern cardinals (Cardinalis cardinalis), and compared these with mate provisioning rates, nestling feeding rates, and nest attentiveness. We found that male ornamentation may not be indicative of mate provisioning rates. Mate provisioning rate did not co‐vary with reproductive success, male feedings to nestlings, or nest attentiveness of females. However, females which were fed more often during incubation tended to provision nestlings less. Reduced female parental effort following extensive incubation feeding may be indicative of females using incubation feeding to assess future male parental effort. Male hormonal condition that favors high rates of nestling provisioning may be a proximate cause of mate provisioning during incubation, even in the absence of selection, favoring high rates of mate provisioning. Both sexes may have capitalized on this unselected behavior.     

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.     

Predation by sparrowhawks Accipiter nisus on male and female pied flycatchers Ficedula hypoleuca in relation to their breeding behaviour and foraging

Bright plumage, song display, and aggressive resource defence in males may cause higher predation on males than on females during the breeding season. However, in birds, higher predation on females is sometimes observed. Parental investment may be high in females (egg-laying, incubation and feeding of offspring), which might lead to a high risk of predation. We studied predation by sparrowhawks Accipiter nisus in relation to behaviour in pied flycatchers Ficedula hypoleuca where breeding males are more conspicuous than females in plumage and behaviour. Male pied flycatchers generally occupied more exposed perches than females. Females were more mobile and foraged more than males, especially prior to and during incubation. During the incubation and nestling stages, when predation on the sexes could be directly compared, sparrowhawks took about the same number of male and female pied flycatchers. During incubation, however, females spent about 77% of the day in the nest and were 4.7 times more vulnerable than males per unit of time available (i.e. outside the nest). A comparison with the chaffinch Fringilla coelebs , where hawks took more females than males, indicates that timing of breeding, foraging behaviour and parental roles of males and females affect predation risk.     

Sensitivity of breeding parameters to food supply in Black-legged Kittiwakes Rissa tridactyla

We fed Herring Clupea pallasi to pairs of Black-legged Kittiwakes Rissa tridactyla throughout the breeding season in two years at a colony in the northern Gulf of Alaska. We measured responses to supplemental feeding in a wide array of breeding parameters to gauge their relative sensitivity to food supply, and thus their potential as indicators of natural foraging conditions. Conventional measures of success (hatching, fledging and overall productivity) were more effective as indicators of food supply than behavioural attributes such as courtship feeding, chick provisioning rates and sibling aggression. However, behaviour such as nest relief during incubation and adult attendance with older chicks were also highly responsive to supplemental food and may be useful for monitoring environmental conditions in studies of shorter duration. On average, the chick-rearing stage contained more sensitive indicators of food availability than prelaying or incubation stages. Overall, rates of hatching and fledging success, and the mean duration of incubation shifts were the most food-sensitive parameters studied.     

Temperature and life history: experimental heating leads female tree swallows to modulate egg temperature and incubation behaviour

1. Life-history decisions are strongly affected by environmental conditions. In birds, incubation is energetically expensive and affected significantly by ambient temperature. We reduced energetic constraints for female tree swallows (Tachycineta bicolor) by experimentally heating nests during incubation by an average of 6.9 degrees C to test for changes in incubation behaviour. 2. Females in heated boxes (hereafter 'heated females') increased time spent incubating and maintained higher on-bout and off-bout egg temperatures. This indicates that female energetic constraints, not maximizing developmental conditions of offspring, determine incubation investment. Furthermore, this result suggests that embryonic developmental conditions in unmanipulated nests are suboptimal. 3. We found individual variation in how females responded to experimental heating. Early-laying (i.e. higher phenotypic quality) females with heated nests increased egg temperatures and maintained incubation constancy, while later-laying (lower quality) heated females increased incubation constancy. Changes in egg temperature were due to changes in female behaviour and not due directly to increases in internal nest-box temperatures. 4. Behaviour during the incubation period affected hatching asynchrony. Decreased variation in egg temperature led to lower levels of hatching asynchrony, which was also generally lower in heated nests. 5. Our study finds strong support for the prediction that intermittent incubators set their incubation investment at levels dictated by energetic constraints. Furthermore, females incubating in heated boxes allocated conserved energy primarily to increased egg temperature and increased incubation attentiveness. These results indicate that studies investigating the role of energetics in driving reproductive investment in intermittent incubators should consider egg temperature and individual variation more explicitly.     

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.     

Nest predation is little affected by parental behaviour and nest site in two African Sylvia warblers

Parental behaviour and nest site are supposed to affect nest predation in birds. Few nest visits and high nest attentiveness are assumed to lead to low predation rates. Poorly concealed nests are thought to be more likely to be preyed upon than well concealed nests. Studies on the relationship between parental behaviour, nest site, and nest predation are rare and none have, so far, been conducted in the Afrotropics. We studied the effect of nest site, nest visitation rate, and nest attentiveness on the nest predation rate of the two tropical warblers Sylvia boehmi and Sylvia lugens in Kenya. Parental behaviour and predation on nests of 13 breeding pairs of both species were observed daily in two consecutive breeding seasons. In both species, parental activity at the nest was low [0.9 trips to the nest in 30 min during incubation, maximum 4.6 (S. boehmi) and 5.8 (S. lugens) trips to the nest in the nestling stage]. Predation rates in both species were high (Mayfield nest success 19.4 and 33.2%). Our analysis revealed only weak evidence for an effect of nest site, nest visitation rate, and nest attentiveness on the predation rate. It is suggested that smaller clutches of tropical in comparison to northern temperate birds result from lower feeding rates in tropical ecosystems with high predation rates (Skutchs hypothesis). The underlying mechanism could not be proven in this study.     

Red-winged blackbirds (Agelaius phoeniceus) with higher baseline glucocorticoids also invest less in incubation and clutch mass

Glucocorticoid hormones facilitate responses to environmental challenges by mediating diverse physiological and behavioral changes, including resource mobilization and altered reproductive effort. Elevated glucocorticoids might indicate that an individual is facing high levels of environmental challenges and thus, elevated concentrations might be associated with reduced fitness (CORT-fitness hypothesis). Alternatively, the energetic demands of reproduction might be a challenge that requires elevated glucocorticoids to mobilize resources to support reproductive effort, ultimately increasing reproductive investment and fitness (CORT-adaptation hypothesis). Investigations of glucocorticoid-fitness relationships have yielded mixed results. Variation in the direction of this relationship could be caused in part by differences in the contexts in which the relationship was assessed. Incorporating context, such as life history stage, could be key to understanding the role of glucocorticoids in influencing fitness outcomes. We investigated the relationship between corticosterone and reproductive effort and success within a single life history stage: incubation of eggs. In an observational study, we measured baseline corticosterone in incubating female red-winged blackbirds (Agelaius phoeniceus), monitored incubation behavior, and determined hatching success for each nest. Incubating birds with higher baseline corticosterone concentrations had more frequent, shorter incubation bouts and spent less time overall incubating their clutches of eggs than birds with lower corticosterone concentrations. Elevated corticosterone was also associated with lower clutch mass, but neither corticosterone nor incubation effort were correlated with hatching success. Although experimental tests are needed to establish causation, these results suggest that during the incubation period, corticosterone might shift resource investment towards self-maintenance, and away from current reproductive effort.     

Mate-sharing costs in polygynous Northern Lapwings Vanellus vanellus

In this study bigamous female Northern Lapwings Vanellus vanellus received significantly less incubation relief from their males than monogamous females. On average, monogamous males spent 34.3% of their time incubating and bigamous males 29.9%. Bigamous males divided their effort between their nests, incubating on average 9.4% on primary nests and 20.5% on secondary nests. Bigamous females compensated for the lack of male relief. Primary females incubated for 71.8% of their time, secondary females for 64.2%, while monogamous females spent 52.7% of their time incubating. As a result, there was no significant difference in total nest attentiveness among nests of different status. Primary and secondary females received equivalent incubation relief from the male. Bigamous males increased their contribution to incubation significantly as the season progressed. A bigamous male's distribution of incubation relief between his females was unrelated to female body mass, or to the degree of asynchrony between primary and secondary females in arrival and laying. Incubation time was significantly, negatively, correlated with total nest attentiveness. Monogamous females spent most time, secondary females spent an intermediate time, and primary females spent the least time on maintenance behaviour (foraging, comfort behaviour, inactivity). No significant differences were found in hatching success among females of different mating status. However, the ratio of unhatched to hatched eggs (i.e. the eggs that remained in the nest at the time of hatching) differed significantly: secondary females hatched a smaller proportion of their eggs than monogamous and primary females.     

Experimental manipulation of female tail length did not cause differential allocation by males in the barn swallow

The evolution and maintenance of female ornamentation has attracted increasing attention, because the previous explanation, that is a non‐functional copy of functional male ornamentation, seems insufficient to explain female ornamentation. A post‐mating sexual selection, differential allocation, may be more common than pre‐mating sexual selection, but few studies have investigated differential allocation by males. Here, we studied differential allocation of incubation investment by male barn swallows Hirundo rustica, a model species for the study of sexual selection, because our previous correlative study demonstrated a positive relationship between female tail length and male incubation investment. We manipulated the length of the outermost tail feathers in females after clutch completion and examined whether males adjust incubation investment according to female ornamentation. Because extra‐pair paternity is virtually absent in the study population, we were able to study differential allocation based on the tradeoff between current and future reproductive investments, rather than the tradeoff between current paternal investment and additional mating effort. The experimental treatment had no significant effect on male nest attentiveness, whereas female tail length before manipulation predicted male nest attentiveness. The observed pattern is consistent with differential access; that is, well‐ornamented individuals have greater access to mates with high reproductive (parental) ability, rather than differential allocation during incubation. Alternatively, males can directly assess eggs in their nests, and thus, as seen in other species, males might adjust their incubation investment based on the egg characteristics of long‐tailed females.     

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

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

The effect of ectoparasite nest load on the breeding biology of the Penduline Tit Remiz pendulinus

In this study we investigated the importance of ectoparasite load in the nest on the breed-ing system of the Penduline Tit Remiz pendulinus, examining the effect of mite abundance in the nest on mate choice, reproductive success and parental effort. The two most common ectoparasites were the Northern Fowl Mite Dermanyssus hirundinis and the Northern Feather Mite Ornithonyssus sylviarum. The results show that mite load is important in mate choice but has no adverse effect on reproductive success. The results also indicate that infestation level is related to the quality of the male (mask-width). Parental feeding rate was negatively related to mite load. This relationship indicated that Penduline Tits did not compensate for higher parasite loads by increasing feeding but rather reflected the con dition of the parent and its investment in self-maintenance behaviour.     

Buildings promote higher incubation temperatures and reduce nest attentiveness in a Neotropical thrush

Incubation is an energetically costly parental task of breeding birds. Incubating parents respond to environmental variation and nest‐site features to adjust the balance between the time spent incubating (i.e. nest attentiveness) and foraging to supply their own needs. Non‐natural nesting substrates such as human buildings impose new environmental contexts that may affect time allocation of incubating birds but this topic remains little studied. Here, we tested whether nesting substrate type (buildings vs. trees) affects the temperature inside the incubation chamber (hereafter ‘nest temperature’) in the Pale‐breasted Thrush Turdus leucomelas, either during ‘day’ (with incubation recesses) or ‘night’ periods (representing uninterrupted female presence at the nest). We also tested whether nesting substrate type affects the incubation time budget using air temperature and the day of the incubation cycle as covariates. Nest temperature, when controlled for microhabitat temperature, was higher at night and in nests in buildings but did not differ between daytime and night for nests in buildings, indicating that buildings partially compensate for incubation recesses by females with regard to nest temperature stability. Females from nests placed in buildings exhibited lower nest attentiveness (the overall percentage of time spent incubating) and had longer bouts off the nest. Higher air temperatures were significantly correlated with shorter bouts on the nest and longer bouts off the nest, but without affecting nest attentiveness. We suggest that the longer bouts off the nest taken by females of nests in buildings is a consequence of higher nest temperatures promoted by man‐made structures around these nests. Use of buildings as nesting substrate may therefore increase parental fitness due to a relaxed incubation budget, and potentially drive the evolution of incubation behaviour in certain urban bird populations.     

Parental investment of male two-spotted goby, Gobiusculus flavescens (Fabricius)

The trade-off between parental care and feeding was studied in the male two-spotted goby (Gobiusculus flavescens F.). Two temperatures, 8.5 degrees C and 13.0 degrees C, were used, with five replicates at each temperature, in order to determine whether temperature influenced parental behaviour. In each replicate, two males and four females were introduced to an aquarium, where the males chose between two nests and courted the females. In each replicate, one male spawned. After spawning, the males guarded the eggs until hatching. The guarding males' behaviour was recorded with a video camera twice a day (15 min each time), once before and once after they were fed. The male's condition (c-factor) was calculated at the start of the experiment and after egg hatching. The eggs were spawned in an artificial nest (half of a PVC tube), and attached to the nest in a single layer. The areas with eggs (representing brood size) were marked after spawning and the fry counted after hatching (which was used to calculate area hatched). Numbers of prey eaten (plankton) and number of aggressive encounters between the guarding male and the other fishes were recorded. Time spent in the nest and time used on fertilisation, fanning and cleaning were estimated and related to egg age, brood size, hatching success, temperature and food availability (no food or food).The results showed that feeding (expected to influence future reproduction) decreased and parental expenditure (current reproduction) increased, as the eggs developed (became closer to independence). Parental expenditure was significantly higher at 13.0 degrees C than at 8.5 degrees C, presumably due to higher oxygen demands by the eggs, and a greater risk of egg-infections. The c-factor of the males guarding eggs decreased over time, in contrast to the non-guarding males' c-factor. Guarding males' aggressiveness decreased as the eggs got older, but increased just before hatching. A possible explanation for this could be the decreasing intrusion by the non-guarding male and females caused by high aggressive behaviour by the guarding male early in the brood cycle. The exploitation of the nest (percentage of total nest area covered by eggs) seemed to determine the amount of parental expenditure and loss of condition, while brood size (area of eggs) had no effect.     

Sex roles, parental experience and reproductive success of eastern kingbirds, Tyrannus tyrannus

We quantified parental behaviour of eastern kingbirds during the incubation and nestling periods to determine parental roles, and to examine the impact of previous breeding experience (defined as having bred on the territory in the past) on behaviour and reproductive success. Females performed all incubation, while males spent more than 60% of their time in vigilant or nest guarding behaviour during incubation. Parental roles were not defined as sharply during the nestling period. Females spent more time vigilant, but males provisioned young at only 54% of the rate of females. Vigilance and nest watching were still primarily male duties. Male and female behaviour did not vary with the pair's combination of experience (e.g. experienced-experienced versus inexperienced-inexperienced in previous-current breeding season, respectively) during either phase of reproduction, but experienced males were more vigilant during incubation and fed young relatively more than inexperienced males. Experienced females were also more efficient foragers. Although behaviour did not differ among the four combinations of pair experience, inexperienced pairs none the less lost the most young to starvation and predation. Consequently, inexperienced pairs fledged one less nestling per nesting attempt than did pairs with at least one experienced breeder. Our results suggest that having at least one experienced breeder substantially improved a pair's reproductive success. We propose that female site fidelity is a safeguard to avoid the lower breeding success a female would incur if she were to move to a new territory and breed with an inexperienced male. Copyright 1999 The Association for the Study of Animal Behaviour.