Male Incubation Feeding Research Articles

Blue tit Cyanistes caeruleus males increase their reproductive effort when subject to a flea experimental manipulation

Parasites exert a strong selection pressure on their hosts as manifested in behavioural antiparasite traits to reduce negative impacts on fitness. The numerous nest‐dwelling ecto‐parasites residing in avian nests make altricial birds excellent model‐systems for investigating the relationship between parasites and their hosts. Here, we experimentally increased natural levels of hen fleas Ceratophyllus gallinae in blue tit Cyanistes caeruleus nests during incubation, and tested their effects on parental incubation behaviours and reproductive performance. Our experimental addition of fleas resulted in an increase in feeding effort of males to incubating females. Frequency of male feedings was also positively related to clutch size. These results suggest that males increase their reproductive effort in flea manipulated and large broods. This will, at least partly, compensate female costs in nests with high ecto‐parasite density and many nestlings. Furthermore, nestling mass at day six in experimental nests decreased with brood size, which was not the case in nests with a natural level of fleas. In line with male incubation feeding, parents may try to compensate for the costs inflicted by the fleas but can only partly compensate when brood size is large.

Patterns of female nest attendance and male feeding throughout the incubation period in Blue Tits Cyanistes caeruleus

Most bird species exhibit biparental care, but the type of care provided by each sex may differ substantially. In particular, during the incubation phase in passerines, females perform most or all of the incubation, while the male cares for the brood indirectly by feeding the female. However, detailed descriptions of this male investment during the incubation period are missing. Here, we quantitatively describe female nest attendance and male incubation feeding throughout the ~ 14‐day incubation period in a population of Eurasian Blue Tits Cyanistes caeruleus breeding in nestboxes. Males and females progressively increased their daily activity at the nest over the incubation period. The amount of day‐time incubation, measured as the proportion of the active day (time interval between first nestbox exit in the morning and last entry in the evening) a female spent inside the nestbox, varied between 52 and 60% with an average of 55% per day. The frequency of male incubation feeding varied between 0 and 74 times per day with an average of 12 feeds per day. Both male feeding rate and female nest attendance were highest in the morning and declined rapidly throughout the day. Females were more likely to be off the nest during the warmest periods (15–21 °C), as expected based on thermal needs of the developing embryos, but also during the coldest periods (2–5 °C), presumably due to the energetic needs of the female. This was despite the fact that males fed their females more often at the nest when ambient temperatures were low. Females that received more feeds incubated more and their off‐nest bouts were shorter after a feed. The observed variation in female incubation and in male feeding rate was not linked to individual age or to variation in measures of reproductive success. However, direct observations showed that in some pairs a substantial amount of feeding by males occurred outside the nestbox. This suggests that the true male investment might have been underestimated, here and in previous studies.

The effect of male incubation feeding on female nest attendance and reproductive performance in a socially monogamous bird

The incubation of eggs plays a key role in avian parental care. To ensure embryo development, incubating parents have to keep their eggs within the appropriate temperature limits. To do so, incubating individuals allocate substantial energy to the thermal demands of their eggs, but they face a trade-off with self-maintenance (own metabolism) because they usually cannot forage while incubating eggs. In species with female-only incubation, males can help their partners by providing them with food on the nest, a behavior which may enable females to spend more time incubating and could, consequently, lead to improved reproductive performance. In the study reported here, we first investigated whether incubation feeding by males affects nest attendance by females in Blue Tits (Cyanistes caeruleus) and subsequently determined how this incubation feeding affects reproductive performance. We found that the female nest attendance tended to increase with increasing amounts of food supplied by their male partner. Thus, males may enable females to incubate more when needed, as suggested by our observation that male incubation feeding was more frequent when the ambient temperature was lower, and especially so when females incubated later in the breeding season (during the study period the ambient temperature decreased rapidly over the breeding season, which is exceptional). Although female nest attendance did not result in a shorter time until the eggs hatched or in higher hatching success, females that attended the nest more produced heavier nestlings. We suggest that the trade-off between self-maintenance and meeting the demands of egg incubation likely tends to be less when females received more assistance from their partner during the egg incubation period, resulting in a higher investment in offspring.

The effect of ambient temperature, habitat quality and individual age on incubation behaviour and incubation feeding in a socially monogamous songbird.

Incubation is an important aspect of avian life history. The behaviour is energetically costly, and investment in incubation strategies within species, like female nest attentiveness and the feeding by the non-incubating partner during incubation, can therefore vary depending on environmental and individual characteristics. However, little is known about the combined effect of these characteristics. We investigated the importance of ambient temperature, habitat quality, and bird age on female incubation behaviour and male feeding of the incubating female (incubation feeding) in blue tits Cyanistes caeruleus, a socially monogamous songbird. An increase in ambient temperature resulted in a higher nest temperature, and this enabled females to increase the time off the nest for self-maintenance activities. Probably as a consequence of this, an increase in ambient temperature was associated with fewer incubation feedings by the male. Moreover, in areas with more food available (more deciduous trees), females had shorter incubation recesses and males fed females less often. Additionally, males fed young females more, presumably to increase such females’ investment in their eggs, which were colder on average (despite the length of recesses and female nest attentiveness being independent of female age). Male age did not affect incubation feeding rate. In conclusion, the patterns of incubation behaviour were related to both environmental and individual characteristics, and male incubation feeding was adjusted to females’ need for food according these characteristics, which can facilitate new insights to the study of avian incubation energetics.Significance statementParents often invest a substantial amount of energy in raising offspring. How much they do so depends on several environmental factors and on the extent they cooperate to raise the offspring. In birds, males can feed incubating females, which may allow females to stay longer on the nest, which, in turn, may ultimately improve reproductive success. The interplay between environmental factors and such incubation feeding on incubation attendance has, however, received little attention. Here, we show that favourable circumstances (higher ambient temperature and food availability) allowed incubating blue tit females to increase the time off the nest to improve self-maintenance and males to feed them less, whereas males also fed inexperienced partners more often. Thus, we show a concerted effect of several environmental and intrinsic factors on parental effort during incubation, which will help to improve the general understanding of avian incubation and parental care.

Males respond to female begging signals of need: a handicapping experiment in the pied flycatcher, Ficedula hypoleuca

The ‘female nutrition’ hypothesis proposes that food provided by males during incubation is an important energy source for females in bird species in which females alone incubate. Females should be able to communicate their needs through begging signals to mates and males may compensate for the energetic limitations of females through their feeding visits, owing to their overlapping reproductive interests. To test whether female begging during incubation is an honest signal of energetic need and whether mates respond to it we experimentally handicapped female pied flycatchers at the beginning of incubation by clipping two primary flight feathers on each wing. Experimental manipulation led females to intensify begging displays arising from condition impairment and males accordingly increased their incubation feeding rates. Female begging intensity explained more than half of the variation in male incubation feeding rate, thereby showing that female nutrition is the main factor explaining male incubation feeding. Moreover, handicapped females consumed a higher proportion of male food deliveries during the first few days after hatching and weighed less at the end of the nestling period than control females. Handicapping had no influence on female incubation behaviour, hatching and breeding success, nestling and male condition or female nestling provisioning. The provisioning rates of males in the late nestling stage were higher in experimental nests. This is the first experimental study showing that males adjust incubation feeding rates to behavioural displays of need by their mates. The ability of females to modify their begging displays according to need may be an important adaptation that allows females to maintain a good energetic condition during incubation.

Females Paired with More Attractive Males Show Reduced Oxidative Damage: Possible Direct Benefits of Mate Choice in Pied Flycatchers

AbstractDirect benefits of female mate choice may concern female fertility and fecundity but also physiological status. In birds with biparental care, males may contribute to improve the condition and health of their pair‐mates through help in constructing nests, incubation or incubation feeding and nestling provisioning. They may also reduce harassment of females by non‐pair males. A consequence of these male activities could be expressed in terms of oxidative damage, which may depend on metabolic effort and social stress. Here, we have related male contribution to parental and territorial duties to female oxidative status in the pied flycatcher Ficedula hypoleuca, a species where preferred males present darker dorsal plumage and, in Iberian populations, a large white forehead patch. Darker males were paired with females with high incubation attendance and reduced nestling provisioning rates, which may lead to reduced female exertion. These males owned nest boxes at which there were fewer visits by non‐pair males. Although females paired with dark mates worked less hard, they were able to raise more fledglings. Female oxidative damage measured as malondialdehyde (MDA) level in plasma declined with increasing incubation attendance and male incubation feeding. Moreover, levels of MDA in females declined with both darkness of male dorsal plumage and male forehead patch size when controlling for female forehead patch size and male age. The effect of male plumage darkness was especially strong. Females paired with middle‐aged males (2–3 yr) showed reduced levels of MDA compared with those paired with 1‐yr‐old and more than 3‐yr‐old males. Male age could not explain the effects of male attractiveness. Females paired with attractive males were more successful in reproduction while suffering reduced oxidative damage, possibly mediated by help during incubation and nestling rearing from their pair‐mates. Although correlative, the evidence suggests direct benefits of females paired with more attractive males.

Predator-induced female behavior in the absence of male incubation feeding: an experimental study

Parental care plasticity is critical to understanding the ecological and evolutionary influence of nest predation on life history strategies. In birds, incubation imposes a trade-off between the requirements of females (i.e., food) and egg requirements (i.e., heat and protection from predators). However, studies on this topic are rare and usually restricted to species where the male feeds the incubating female, relaxing her incubation costs. Males and females can reduce their activity at the nest to avoid detection by predators. However, females could follow two alternative antipredator strategies: to delay their return to the nest to avoid attracting attention from the potential predator or to return to the nest as soon as possible to enhance nest concealment. In this study, we manipulated the perceived risk of nest predation of incubating common blackbirds (Turdus merula), a species without incubation feeding, to study female behavioral changes induced by nest predation risk. We show experimentally that female blackbirds can reduce their nest visits in the situation with higher nest predation risk. In addition, we confirm that females significantly delay their return to the nest in the presence of a nest predator, contradicting the nest concealment hypothesis. However, our results could be interpreted as a passive antipredator response (to minimize clues given to predators) or as an active antipredator response (to search for predators to expel them from their territories).

Male incubation feeding in songbirds responds differently to nest predation risk across hemispheres

Evolution of parental care behaviour has been of considerable interest to behavioural ecologists for a long time. Incubation feeding, where an individual incubating eggs is provisioned by another individual, is an important component of avian parental care. It may be critical for breeding success by allowing the incubating bird to spend more time on the eggs. However, very little is known about environmental factors shaping incubation feeding, and incubation behaviour in general, of tropical and southern hemisphere birds, and how this differs compared to northern hemisphere species. We collated available data on the rate of incubation feeding in Australian, New Zealand and North American songbirds (78 species from 25 families). There was a strong positive relationship between female incubation attentiveness and incubation feeding by males; however, female attentiveness was higher in North America than in Australia and New Zealand for the same intensity of male incubation feeding. Incubation feeding was not related to species body mass, social organization, geographical latitude or ambient temperature. It differed significantly between families, but overall was not different between regions. Incubation feeding rate was related to nest predation rate, but differently in the two regions. It increased with nest predation rate in Australia and New Zealand, but decreased with nest predation rate in North America. We suggest that this may be caused by different predator–prey interactions in North America versus Australia and New Zealand, which could have shaped the evolution of incubation feeding differently. ► We model rates of male incubation feeding in songbirds across two continents. ► We use data obtained by a comprehensive review of published literature. ► Incubation feeding decreased with nest predation in North America. ► Incubation feeding increased with nest predation in Australia and New Zealand. ► Nest attentiveness increased with incubation feeding in both continents.

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

AbstractParental 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.

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

AbstractBecause 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.

Effects of ambient temperature on avian incubation behavior

Ambient temperature is commonly thought to influence avian incubation behavior. However, results of empirical studies examining correlations between ambient temperature and bout duration are equivocal. We propose that these equivocal results can be partly explained by developing a conceptual understanding of how we should expect temperature to influence incubation. We demonstrate why linear correlation analyses across a wide range of temperatures can be inappropriate based on development of an incubation model for small birds that incorporates how ambient temperature influences both embryonic development and adult metabolism. We found support for predictions of the model using incubation data from orange-crowned warblers (Vermivora celata) in Arizona. Both off- and on-bout duration were positively correlated with ambient temperature between 9 and 26C, but unrelated to ambient temperature 9 and 26–40C. Bout durations declined as ambient temperature approached or exceeded 40C. Incubating orange-crowned warblers appeared to avoid bouts off the nest 7 min and bouts on the nest 20 min. Time of day, duration of the previous bout, and variation among nests all explained variation in both onand off-bout duration. Although we found support for the general shape of the incubation model, temperature still explained only a small portion of the overall variation in on- and off-bout duration. Results of previous studies were generally consistent with the model for off-bout duration; most studies in colder environments reported positive correlations with temperature, and the one negative correlation reported was from a hot environment. However, the relationships between on-bout duration and temperature reported in previous studies were less consistent with our model and our data. Although some discrepancies could be explained by considering our model, some studies reported negative correlations in cold environments. The effect of ambient temperature on duration of on-bouts probably differs among species based on the amount of fat reserves females typically carry during incubation and the extent of male incubation feeding. Additional studies of the effects of temperature on avian incubation will help improve the general model and ultimately aid our understanding of energetic and ecological constraints on avian incubation. Key words: ambient temperature, foraging, incubation behavior, incubation model, incubation rhythm, nest attentiveness, on-bout duration, off-bout duration. [Behav Ecol 11:178–188 (2000)]

Clutch Size Manipulation in the Pied Flycatcher: Effects on Nestling Growth, Parental Care and Moult

To study parental responses during the incubation and nestling periods, nestling growth and adult moult, two eggs were either removed from or added to Pied Flycatcher Ficedula hypoleuca clutches shortly after clutch completion. Incubation period and hatching success did not differ between experimental groups. In the enlarged group the rate of male incubation feeding of the female was higher than in the other groups. Fledgling mass and tarsus length were negatively related to the experimental change in clutch size, and this effect arose entirely in the first four days after hatching. The results suggest that the embryo development during the incubation period and nestling growth during the first days after hatching are important for the subsequent nestling growth and should be taken into account when analysing the results of brood size manipulations. Fledging and breeding success, fledgling mass and tarsus length differed between experimental groups with lower values in the enlarged group. There was an effect of experimental treatment on female and male feeding rates, with higher number of feeding visits in the enlarged group. The onset of adult moult was related to calendar date and there was no effect of the experiment on the proportion of birds moulting at any given time. At least in this population, moult may not be a mechanism mediating costs of reproduction.

Female nutritional state affects the rate of male incubation feeding in the pied flycatcher Ficedula hypoleuca

Male pied flycatches Ficedula hypoleuca regularly feed their mates during incubation. By experimentally supplying some females with extra food we studied how the female's nutritional state affected her incubation schedule and the rate at which her mate fed her. Females that received extra food spent more time on the nest and shorter periods away from it, compared with control females. This suggests that nest attentiveness is governed by the amount of energy available to the female. When females reccived extra food, males decreased their rate of incubation feeding. They also did so in response to increasing ambient temperatures, whereas incubation schedules were unaffected. We, therefore, conclude that our results support the “female nutrition hypothesis”, i.e., that the food provided by the male constitutes a significant nutritional contribution to the incubating female.

Allocation of incubation feeding in a polygynous mating system: a study on pied flycatchers Ficedula hypoleuca

The way in which male pied flycatchers Ficedula hypoleuca allocated their incubation feeding efforts to their mates was studied. The overall rate of male incubation feeding was negatively correlated with the ambient temperature and positively correlated with the physical condition of the males. Polygynously mated females received significantly fewer feeds than did monogamously mated females. No difference in the food delivery rates to primary and secondary females was found. The pattern of investment by polygynous males was unrelated to the time-lag between completion of the different clutches and to any difference in clutch size. The reduction in the delivery rate to polygynously mated females had a negative effect on their incubation efficiency, since they spent a greater proportion of the time off the nest and the duration of the incubation period was prolonged compared to that of monogamously mated females. Male feeding behaviour is discussed in relation to the reproductive value represented by the nests and to the costs and benefits of incubation feeding.