Females compensate for moult‐associated male nest desertion in Hooded Warblers

Uniparental offspring desertion occurs in a wide variety of avian taxa and usually reflects sexual conflict over parental care. In many species, desertion yields immediate reproductive benefits for deserters if they can re‐mate and breed again during the same nesting season; in such cases desertion may be selectively advantageous even if it significantly reduces the fitness of the current brood. However, in many other species, parents desert late‐season offspring when opportunities to re‐nest are absent. In these cases, any reproductive benefits of desertion are delayed, and desertion is unlikely to be advantageous unless the deserted parent can compensate for the loss of its partner and minimize costs to the current brood. We tested this parental compensation hypothesis in Hooded Warblers Setophaga citrina, a species in which males regularly desert late‐season nestlings and fledglings during moult. Females from deserted nests effectively doubled their provisioning efforts, and nestlings from deserted nests received just as much food, gained mass at the same rate, and were no more likely to die from either complete nest predation or brood reduction as young from biparental nests. The female provisioning response, however, was significantly related to nestling age; females undercompensated for male desertion when the nestlings were young, but overcompensated as nestlings approached fledging age, probably because of time constraints that brooding imposed on females with young nestlings. Overall, our results indicate that female Hooded Warblers completely compensate for male moult‐associated nest desertion, and that deserting males pay no reproductive cost for desertion, at least up to the point of fledging. Along with other studies, our findings support the general conclusion that late‐season offspring desertion is likely to evolve only when parental compensation by the deserted partner can minimize costs to the current brood.     

Mate desertion by male Great Reed Warblers Acrocephalus arundinaceus at the end of the breeding season

Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains. Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains.     

Factors affecting the cost of polygynous breeding for female Great Reed Warblers Acrocephalus arundinaceus

The male's role in feeding young and his effect on the mortality of eggs and nestlings of the Great Reed Warbler Acrocephalus arundinaceus were studied in Kahokugata, central Japan, during eight breeding seasons. Three potential costs of polygynous breeding for females were examined:(i) a reduction of male parental care per brood, (ii) an attraction of predators due to concentration of nests, and (iii) an increase of unfertilized eggs due to the reduced frequency of copulation per female. Only the first cost was found to occur:polygynous males fed their young, especially of their second broods, less frequently than monogamous ones. The lowered paternal feeding frequency, however, did not increase the mortality by starvation in second-hatched broods. This was because the warm climate emancipated females from brooding and enabled them to compensate for the deficiency of feeding by males. The paper discusses possible roles of the staggering of the time of laying among harem mates and the shift of breeding status of females caused by nesting failure, in reducing potential costs of polygynous breeding for females.     

Mate desertion by male Great Reed Warblers Acrocephalus arundinaceus at the end of the breeding season

Male Great Reed Warblers usually take part in the care of offspring as nest defenders and by feeding young, but at the end of the breeding season they desert their mates with eggs or nestlings. Deserted females continue offspring care. Desertion does not depend on the male's mated status (polygynous or monogamous) nor on his past breeding success. Deserted females compensate for the loss of their partners by increasing the frequency of food-bringing, resulting in a reduction in the amount of time the nestlings are brooded. Although desertion may lead to increased rates of offspring mortality through predation, breeding success of deserted females was high, especially if the male assisted during the early stages. Deserters pay costs by giving up the chance of additional matings and by lowering the reproductive success of existing mates. Male warblers reduce the former cost by choosing the season of desertion and the latter is lowered by the female's high parental ability. A deserter was found to start moulting while his mate was still feeding his nestlings, and an earlier start to the moult may be the primary benefit that he gains.     

Does Social Mating System Influence Nest Defence Behaviour in Great Reed Warbler (Acrocephalus arundinaceus) Males?

In birds with biparental care, males and females often conflict over how much care to provide to their offspring and it may be substantially influenced by increased level of polygamy. In accordance with sexual conflict theory, males of socially polygynous bird species provide much less care to their nestlings than do males of most socially monogamous species. Most of previous studies, however, have used feeding behaviour as an index for variations in male parental care only. However, this may be skewed if polygynous males compensate for lower feeding assistance through the provision of other parental care such as protection of nests from predators. In this paper, we examine nest defence behaviour in the facultatively polygynous great reed warbler with respect to sex and type of social mating system. We recorded latency to the first arrival, distance from the predator and defensive reaction of each parent towards a human intruder. Socially polygynous males with two simultaneously active nests defended primary females’ nests less vigorously than socially monogamous males, whereas no differences were found between monogamous and primary females. Generally, however, they took a bigger role in nest defence than males in all cases. Our results support an idea that sexual conflict is driven by polygamy and that type of social mating system can influence nest defence behaviour of facultatively polygynous birds. This finding should be taken into consideration when studying nest defence parental care in polygynous mating systems.     

Male parental care promotes early fledging in an open-nester, the Willow Warbler Phylloscopus trochilus

The importance of male parental care to female reproductive success was investigated in the monogamous Willow Warbler Phylloscopus trochilus by removing the male parent at two different stages of the breeding cycle. Females that were widowed at the start of egg-laying continued breeding and managed to raise their brood on their own with no apparent reductions in numbers fledged or fledgling body-mass. The widowed females compensated for the loss of male assistance by increasing their own food provisioning rate as compared with control females. However, widows spent less time brooding the small young, and the growth rate of nestlings was reduced. In nests where the male parent was removed 7 days after the eggs hatched, the subsequent growth rate of nestlings was still affected, which suggests that male care is influential throughout the nestling period. On average, broods reared by widows fledged 2 days later than did broods of control females. An extension of the nestling period may appreciably affect reproductive success, since 68% of nests failed due to predation, mostly during the nestling period. We suggest that the main role of male parental care in the Willow Warbler is to assure a high growth rate of nestlings, which leads to early fledging and hence a reduced risk of nest predation.     

Parental care and social mating system in the Lesser Spotted Woodpecker Dendrocopos minor

The sexes’ share in parental care and the social mating system in a marked population of the single‐brooded Lesser Spotted Woodpecker Dendrocopos minor were studied in 17 woodpecker territories in southern Sweden during 10 years. The birds showed a very strong mate fidelity between years; the divorce rate was 3.4%. In monogamous pairs, the male provided more parental care than the female. The male did most of the nest building and all incubation and brooding at night. Daytime incubation and brooding were shared equally by the sexes, and biparental care at these early breeding stages is probably necessary for successful breeding. In 42% of the nests, however, though still alive the female deserted the brood the last week of the nestling period, whereas the male invariably fed until fledging and fully compensated for the absent female. Post‐fledging care could not be quantified, but was likely shared by both parents. Females who ceased feeding at the late nestling stage resumed care after fledging. We argue that the high premium on breeding with the same mate for consecutive years and the overall lower survival of females have shaped this male‐biased organisation of parental care. In the six years with best data, most social matings were monogamous, but 8.5% of the females (N=59) exhibited simultaneous multi‐nest (classical) polyandry and 2.9% of the males (N=68) exhibited multi‐nest polygyny. Polyandrous females raised 39% more young than monogamous pairs. These females invested equal amounts of parental care at all their nests, but their investment at each nest was lower than that of monogamous females. The polyandrously mated males fully compensated for this lower female investment. Polygynous males invested mainly in their primary nest and appeared to be less successful than polyandrous females. Polyandry and polygyny occurred only when the population sex ratio was biased, and due to strong intra‐sexual competition this is likely a prerequisite for polygamous mating in Lesser Spotted Woodpeckers.     

Effect of altitude on male parental expenditure in Mountain Bluebirds ( Sialia currucoides ): are higher-altitude males more attentive fathers?

Male investment of time and energy in caring for offspring varies substantially both between and within bird species. Explaining this variation is of long-standing interest to ornithologists. One factor that may affect male care is breeding site altitude, through its effects on climate. The harsher, less predictable abiotic conditions at higher altitudes are hypothesized to favour increased male investment of time and energy in offspring care. We tested this hypothesis by comparing male parental behaviour in Mountain Bluebirds (Sialia currucoides) nesting at 1500 and 2500 m a.s.l. in the Bighorn Mountains of Wyoming, USA. We compared rates of prey delivery to nestlings at these two altitudes at two times: 1–2 days after hatching, when females spend much of their time brooding young, and 12–13 days later, when brooding has ended and nestling energy demands are peaking. High-altitude males fed nestlings 18 and 28% more often than low-altitude males early and later in the nestling stage, respectively, but only the difference in late-stage feeding rates were significant. Like males, females at the high site also fed nestlings significantly more often than females at the low site later in the nestling stage (45% difference in feeding rates). Consequently, the proportion of all feeding trips made by males at the high site (40%) did not differ significantly from that at the low site (44%). Parents at the high altitude may feed nestlings more often to compensate for their greater thermoregulatory costs. Parents may also be attempting to assist nestlings in storing fat and/or attaining a large size and effective homeothermy as quickly as possible to enhance nestling ability to survive bouts of severe weather which are common at high altitudes.     

Who Cares? Males Provide Most Parental Care in a Monogamous Nesting Cuckoo

Cuckoos hold a prominent position in the study of parental care, because they show the greatest variation of any bird family in the way they care for their offspring. Despite this, few data are available on cuckoos with biparental care even though it is the most common form of parental care in cuckoos and birds in general. Here I describe the breeding behaviour of the pheasant coucal, Centropus phasianinus , an old-world nesting cuckoo. I show that males almost exclusively build the nest and incubate and brood the young alone both at night and during the day. The incubation pattern of short, c . 40 min bouts on the nest interspersed with 20 min frequent recesses is well suited to incubation by a single parent and is widespread amongst birds. Males also defend the nestlings and deliver 80% of the feeds to the young consisting of insects (65%), frogs and reptiles. Females did not compensate for their fewer feeding visits by delivering more vertebrates than males. Although coucal males get little help feeding, the hourly feeding rate of 3.8 feeds per brood (1.4 feeds/nestling) exceed that of the few nesting cuckoos studied to date and matches that of other tropical non-passerines. Predominantly male care is found in less than 5% of birds, mainly species with reversed sexual size dimorphism or reversed sex-roles. Its evolution, especially in monogamous species, remains puzzling. The breeding behaviour of pheasant coucals illustrates a possible transitional stage in the evolution of polyandry and interspecific brood parasitism in cuckoos.     

Why do pied flycatcher females mate with already-mated males?

The contribution of the pied flycatcher Ficedula hypoleuca male in parental care was studied to examine why some females mate with already-mated males. No difference in feeding rate was found between older and yearling monogamous males, when comparing nests at the same time. Monogamous and primary females were helped significantly more in parental care by the male than were secondary females of polygynous males. Females could only partly compensate for the absence of a male and of nestlings were reduced in nests with low male assistance. Differences in mate and territory quality were far too slight to make it advantageous for females to choose already-mated males instead of mating with monogamous males. We suggest that males, by being polyterritorial, deceive females into accepting polygyny; and females can be deceived since they do not have time to find out the marital status of males.     

Intraseasonal effects of clutch manipulation on parental provisioning and residual reproductive value of eastern phoebes (Sayornis phoebe)

Summary First clutches of double-brooded eastern phoebes Sayornis phoebe were manipulated (up two eggs, down 2 eggs or no change) to test for intraseasonal reproductive tradeoffs and to test whether size of first brood influenced food delivery rates to nestlings and nestling quality in second broods.Considering all nests from both broods, rate of feeding nestlings increased linearly with brood size but nestling mass per nest decreased with increasing brood size. High nestling weights in small broods may have resulted from parents delivering better quality food, but we did not test this.Among treatment groups in first broods, nestlings from decreased broods weighed more than those in control or increased broods. Treatment did not influence the likelihood that second nests would be attempted after successful first nests nor did it alter the interval between nests. Nestlings of parents that renested weighed more than those of parents that did not, regardless of treatment, suggesting that post-fledging care may preclude renesting. Mass of individual females did not change between broods, regardless of brood size. Clutch sizes of second attempts were not affected by manipulations of first broods but increasing first broods reduced the number of nestlings parents were able to raise to day 11 in their second broods. However, manipulation of first broods did not affect mean nestling mass per nest of nestlings that survived to day 11.In phoebes, parents of small first broods are able to raise nestlings in better condition. We predict that in harsh years, parents of small first broods would be more likely to renest. Parents of enlarged first broods sacrificed quality of offspring in second broods, which seems a reasonable strategy if nestlings from second broods have lower reproductive value.     

Hatching asynchrony and brood reduction influence immune response in Common Kestrel Falco tinnunculus nestlings

The onset of incubation before the end of laying imposes asynchrony at hatching and, therefore, a size hierarchy in the brood. It has been argued that hatching asynchrony might be a strategy to improve reproductive output in terms of quality or quantity of offspring. However, little is known about the mediating effect of hatching asynchrony on offspring quality when brood reduction occurs. Here, we investigate the relationship between phenotypic quality and hatching asynchrony in Common Kestrel Falco tinnunculus nestlings in Spain. Hatching asynchrony did not increase breeding success or nestling quality. Furthermore, hatching asynchrony and brood reduction had different effects on nestlings’ phytohaematogglutinin (PHA)‐mediated immune response and nestling growth. In asynchronous and reduced broods (in which at least one nestling died), nestlings showed a stronger PHA‐mediated immune response and tended to have a smaller body size compared with nestlings raised in synchronous and reduced broods. When brood reduction occurred in broods hatched synchronously, there was no effect on nestling size, but nestlings had a relatively poor PHA‐mediated immune response compared with nestlings raised in asynchronous and reduced broods. We suggest that resources for growth can be directed to immune function only in asynchronously hatched broods, resulting in improved nestling quality, as suggested by their immune response. We also found that males produced a greater PHA‐mediated immune response than females only in brood‐reduced nests without any effect on nestling size or condition, suggesting that females may trade off immune activities and body condition, size or weight. Overall, our results suggest that hatching pattern and brood reduction may mediate resource allocation to different fitness traits. They also highlight that the resolution of immune‐related trade‐offs when brood reduction occurs may differ between male and female nestlings.     

Brood desertion and polygamous breeding in the Kentish Plover Charadrius alexandrinus

In the Kentish Plover Charadrius alexandrinus one of the adults, typically the female, deserts the brood when the chicks are a few days old. Once parental care is terminated, adults may initiate a second nesting attempt if sufficient time remains within the season. For these nests, individuals pair with different mates from those of the first nesting attempt, thus becoming sequentially polygamous. In a small population of Kentish Plovers in Fuente de Piedra lake (southern Spain), the duration of biparental care of broods was longer than in other localities. It also showed considerable variation between years that was evidently related to Gull-billed Tern Gelochelidon nilotica predation pressure on the chicks. There was year-to-year variation in the number of polygamous matings. Both the duration of the breeding season and nesting success in the first half of the season limited the occurrence of polygamy. Despite females deserting broods earlier than males, the interval between the first and second nesting of polyandrous females and polygynous males was similar. The interval was not affected by the body condition of females after the first nesting attempt, nor by problems related to egg formation ability, but was probably due to the availability of potential mates. More females than males initiated second nests, suggesting that polygamous opportunities were more limited for males than for females. In terms of delayed breeding, reduced survivorship or reduced breeding opportunities in years following polygamous breeding, polygamous individuals did not have greater costs than non-polygamous ones. Females with second nests did not seem to be selective in mate choice, mating with any available male. Mates for second nests may therefore be of lower quality than those for first nests, as judged by male plumage characteristics. Clutch sizes and egg characteristics of polyandrous females were similar in first and second nests. Nest success of second nests was only 40% of that of first ones, with nest desertion accounting for 60% of the losses. As the costs of polygamy are apparently low and as breeding success is very variable among years, polygamous breeding of the long-lived Kentish Plover may be an important breeding strategy with which to increase individual lifetime reproductive success.     

Ecological constraints on breeding system evolution: the influence of habitat on brood desertion in Kentish plover

1. One of the fundamental insights of behavioural ecology is that resources influence breeding systems. For instance, when food resources are plenty, one parent is able to care for the young on its own, so that the other parent can desert and became polygamous. We investigated this hypothesis in the context of classical polyandry when females may have several mates within a single breeding season, and parental duties are carried out largely by the male. 2. We studied a precocial wader, the Kentish plover Charadrius alexandrinus, that exhibits variable brood care such that the chicks may be raised by both parents, only by the female or, more often, only by the male. The timing of female desertion varies: some females desert their brood at hatching of the eggs and lay a clutch for a new mate, whereas other females stay with their brood until the chicks fledge. Kentish plovers are excellent organisms with which to study breeding system evolution, as some of their close relatives exhibit classical polyandry (Eurasian dotterel Eudromias morinellus, mountain plover Charadrius montanus), whereas others are polygynous (northern lapwing Vanellus vanellus). 3. Kentish plovers raised their broods in two habitats in our study site in southern Turkey: saltmarsh and lakeshore. Food intake was higher on the lakeshore than in the saltmarsh as judged from feeding behaviour of chicks and adults. As the season proceeded and the saltmarsh dried out, the broods moved toward the lakeshore. 4. As the density of plovers increased on lakeshore, the parents spent more time defending their young, and female parents stayed with their brood longer on the lakeshore. 5. We conclude that the influence of food abundance on breeding systems is more complex than currently anticipated. Abundant food resources appear to have profound implications on spatial distribution of broods, and the social interactions between broods constrain female desertion and polyandry.     

Parental Sex Differences in Food Allocation to Junior Brood Members as Mediated by Prey Size

Individual offspring within a brood may receive different amounts of provisioning from the male and female parents. Some hypotheses suggest that this bias is the result of an active and adaptive choice by parents. An alternative hypothesis is that feeding biases arise as a result of a constraint of fitting large prey items into small gapes. In an experiment with pied flycatchers, Ficedula hypoleuca , we tested for sex-biased allocation to junior nestlings in asynchronous broods and whether this could be explained by active parental choice or by passive allocation according to prey size and gape size. In both control broods and broods with experimentally increased degree of asynchrony, prey types did not differ between parents but females brought smaller prey than males at younger but not older nestling stages. At younger but not older nestling stages, the majority of feeds to junior nestlings were from females, and the smaller nestlings consumed smaller prey than older siblings. However, there was no evidence of active preference of small nestlings by females as parents did not differ in the tendency to bypass a begging senior nestling in order to feed a junior nestling. Provisioning rates by females were lower than those by males when nestlings were young and we suggest that foraging time constraints caused by the need to brood offspring result in females bringing smaller prey than males. In turn, the larger prey brought by males was more often transferred to larger offspring after the smaller ones failed to swallow it. In such cases, 'preferential' feeding of small nestlings by females may simply be a passive side effect of foraging constraints and gape-size limitations.     

Parental care behavior in the monogamous,sexually dimorphic Madagascar paradise flycatcher: sex differences and the effect of brood size

I studied the parental care behavior of the Madagascar paradise flycatcher Terpsiphone mutata in northwestern Madagascar. I especially focused on feeding, brooding and vigilance behaviors. Feeding rate did not differ between males and females, but females spent more time at the nest than males. Females dedicated their time to brooding, while males perched on the nest and were vigilant. Both parents changed the feeding rate in relation to brood size, so the feeding rate per nestling was not different among nests of different brood size. Duration of brooding by females increased with decreasing brood size, suggesting that the Royama effect, the pattern of lower feeding rate per nestling in larger broods, did not apply in this study. Males spent more time on vigilance than females. Anti-predator vigilance by males should be important for nestling survival given the high predation pressure typical of this population. In conclusion, males provide considerable parental care probably to minimize nestling starvation and to avoid nest predation. My results are not consistent with the general pattern of less parental effort by males in monogamous, sexually dimorphic species.     

Brood desertion in Kentish Plover Charadrius alexandrinus: an experimental test of parental quality and remating opportunities

Uniparental male care combined with polyandry is rare in birds, and the best known examples are in shorebirds Charadrii. There are two current hypotheses explaining why males care for the brood, whereas females desert and remate: either males are more capable than females at providing uniparental care (“parental quality hypothesis”) or females gain a greater increase in reproductive success by deserting than do males (“remating opportunity hypothesis”). I experimentally tested both hypotheses in Kentish Plover Charadrius alexandrinus, one of the few avian species in which either parent may desert the brood. By experimentally removing one parent when the chicks hatched, I found that male-tended broods had better survival than female-tended ones, particularly up to 6 days after hatching. It is unlikely that differential brood mortality was caused by chilling of the chicks, since the brooding behaviour of males and females was not different. The results of this study are consistent with the explanation that male-tended broods survived better because males were better able to protect the brood from attacks by conspecifics and predators. The remating opportunity hypothesis was also corroborated because single females acquired new mates faster than did single males. The results of this study suggest that both the better parental capability of males and the greater remating opportunities of females predispose Kentish Plovers for uniparental male care, desertion by the female parent and sequential polyandry.     

Brood provisioning rates and fledgling behavior of Cordilleran Flycatchers in southwestern Colorado

The behavior of young songbirds after fledging is one of the least understood phases of the breeding cycle, although parental provisioning rates and movement of fledglings are key to understanding life history evolution. We studied Cordilleran Flycatchers (Empidonax occidentalis) at two sites in southwestern Colorado, USA, from 2012 to 2017. We banded and sexed breeding adults to determine the relative contributions of males and females to nestling and fledgling care, and attached radio‐transmitters to nestlings to facilitate observations of brood behavior after fledging. Females made 60% and 78% of total observed feedings of nestlings and fledglings, respectively. Parental provisioning rates increased with nestling age, and per‐nestling provisioning rates increased with brood size. Parental provisioning rates declined just before fledging, then increased just after fledging. Fledging times of individuals in broods were asynchronous and concentrated during the late afternoon and early evening. Males stopped caring for fledglings before females even though this species is single‐brooded, with some late‐season broods being abandoned by males. Broods spent the first three weeks after fledging within 400 m of nests, after which they began to disperse. Most aspects of the breeding biology of Cordilleran Flycatchers in our study, including the duration of nestling and fledging periods, female‐dominated provisioning, and movement patterns of fledglings, were similar to those of other Empidonax species. However, the times when young fledged were not concentrated in the morning as reported in most other songbirds, and this result warrants additional study of the timing of fledging in ecologically and taxonomically similar species. The increased per‐nestling provisioning rate with increasing brood size was unexpected, and additional study is needed to determine if this increase results from a trade‐off between adult annual survival and productivity favoring increased provisioning of young in larger broods, or from the existence of high‐quality individuals where larger clutches and higher provisioning rates are linked.     

Food availability and the male's role in parental care in double-brooded Treecreepers Certhia familiaris

The aim of this work was to examine differences in paternal and maternal care in a double-brooded, monogamous species, the Treecreeper Certhia familiaris, in relation to food availability. As a measure of parental care, we recorded the hourly feeding activity of parents when the nestlings from their first and second breeding attempts were 7 and 12 days old. Feeding frequency of the first brood increased with the age of the nestlings and also with the brood size when 12 days old. While the feeding activities of the females were similar with respect to the first and second broods, the males were less active and failed to provide any food to their nestlings in 15 cases out of 28 second broods. In spite of this, the fledglings from the second broods were heavier than those in the first. Such a pattern of male behaviour was possible without being a disadvantage to the chicks because the food supply increased during the breeding season and the female could provide food for the young alone. Thus paternal care was particularly important in times of poor food supply, i.e. during the first brood, where the extent of these males' activity in feeding the 7-day-old nestlings was positively correlated with the average mass of the nestlings. Our results support the idea that the male of monogamous, altricial bird species often makes important contributions to raising the young, especially during periods when it is difficult for the female to do so alone. Males show flexibility in their pattern of parental care, and male Treecreepers change their contribution to the first and second broods within the same season.