Relative influence of male and female care in determining nestling mass in a migratory songbird

Biparental care is common in birds, with the allocation of effort being highly variable between the sexes. In most songbird species, the female typically provides the most care early in the breeding cycle with both parents providing care when provisioning young. Food provisioning should be directly related to offspring quality; however, the relative influence each parent has on offspring quality has rarely been assessed at the nest level. Consequently, we were interested in assessing the relative influence male and female provisioning has on one measurement of offspring quality, nestling mass, in the black‐throated blue warbler Dendroica caerulescens. Over a six year period, 2003–2008, we collected information on average nestling mass per brood on day 6 of the nestling cycle and parental provisioning rates on day 7 of the nestling cycle from 182 first brood nests on three different study plots. We found that average nestling mass was directly related to male provisioning rate, while it was not related to female provisioning rate. On the other hand, estimated biomass provisioned had little influence on average nestling mass, calling into question its utility in assessing parental quality. Finally, there was some indication that parental influence on average nestling mass was dependent on the other parent's provisioning rate, suggesting that parents work in concert to influence nestling quality.     

Postfledging parental care in Savannah sparrows: sex, size and survival

We investigated postfledging parental care in a philopatric population of Savannah sparrows,Passerculus sandwichensis , breeding on Kent Island, New Brunswick, Canada in an effort to understand the factors influencing adult birds' decisions about parental investment in offspring. Brood division was not based on offspring sex: male and female parents were equally likely to care for sons or daughters. The total duration of parental care, from hatching to independence, was similar for sons and daughters (median=23 days), regardless of the sex of the care-giving parent. The duration of parental care also corresponded closely to the time required for juveniles to acquire basic foraging skills. Despite high levels of extrapair paternity, male Savannah sparrows invested as much in postfledging care and were as effective as females in caring for fledglings, based on recruitment of fledglings into the breeding population the following year. Male parents were more likely to care for smaller fledglings and for offspring from early broods (presumably to enable females to dedicate their efforts towards second clutches). Caring for fledglings was costly for parents: survivorship decreased as a function of the duration of postfledging parental care and the number of fledglings cared for. Parental survivorship, however, was not affected by the sex of the fledglings cared for. This study suggests that sex-biased provisioning may be unlikely except in species with strongly sexually dimorphic offspring, biased offspring sex ratios and sex-biased natal dispersal. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

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.     

Costs of egg-laying and offspring provisioning: multifaceted parental investment in a digger wasp

Nest-building Hymenoptera have been a major testing ground for theories of parental investment and sex allocation. Investment has usually been estimated by the likely costs of offspring provisioning, ignoring other aspects of parental care. Using three experimental treatments, we estimated the costs of egg-laying and provisioning separately under field conditions in a digger wasp Ammophila pubescens. In one treatment, we increased the provisioning effort required per offspring by removing alternate prey items as they were brought to the nest. In two other treatments, we reduced parental effort by either preventing females from provisioning alternate nests or preventing them from both ovipositing and provisioning. Our results indicate that both egg-laying and provisioning represent significant costs of reproduction, expressed as differences in productivity but not survival. A trade-off-based model suggests that other components of parental care such as nest initiation may also represent significant costs. Costs of egg production and nest initiation are probably similar for male and female offspring, so that taking them into account leads to a less male-biased expected sex ratio. Mothers compensated only partially for prey removal in terms of the total provisions they gave to individual offspring.     

Provisioning in western bluebirds is not related to offspring sex

Parents should invest more in one sex of offspring if the fitnessreturn per unit investment is higher for that sex. Sex-biasedprovisioning may occur when sons and daughters differ in theirneeds or when there is local resource competition or enhancement.We used feeding observations and sex-ratio manipulations todetermine if sex-biased provisioning occurs in western bluebirds(Sialia mexicana). The sex ratio of the brood had no effecton feeding rates by adults at unmanipulated nests over a 6-yearperiod. Adult males and females also did not differ in the numberof feedings made to sons and daughters in 13 videotaped nests.Likewise, adult feeding rates to nests experimentally biasedtoward sons or daughters did not differ significantly. Nesdingsthat were closest to the nest hole and that reached highestwere the most likely to be fed. Sons and daughters did not differin the begging behaviors most likely to result in a feeding.We conclude that sex-biased provisioning does not occur in thispopulation of western bluebirds and diat nesding behavior maybe a more important determinant of feeding.     

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.     

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.     

Pair bond duration influences paternal provisioning and the primary sex ratio of brown thornbill broods

Parents should vary their level of investment in sons and daughters in response to the fitness costs and benefits accrued through male and female offspring. I investigated brood sex ratio biases and parental provisioning behaviour in the brown thornbill, Acanthiza pusilla, a sexually dimorphic Australian passserine. Parents delivered more food to male-biased than female-biased broods. However, factors determining parental provisioning rates differed between the sexes. Female provisioning rates were related to brood sex ratio in both natural and experimental broods with manipulated sex ratios. In contrast, male provisioning rates were not affected by brood sex ratio in either natural or experimental broods. However, males in established pairs provisioned at a higher rate than males in new pairs. Data on the sex ratio of 109 broods suggest that female brown thornbills adjust their primary sex ratio in response to pair bond duration. Females in new pairs produced broods with significantly fewer sons than females in established pairs. This pattern would be beneficial to females if the costs of rearing sons were higher for females in new than established pairs. This may be the case since females in new pairs provisioned experimental all-male broods at elevated rates. The condition of nestlings also tended to decline more in these all-male broods than in other experimental broods. This will have additional fitness consequences because nestling mass influences recruitment in thornbills. Female thornbills may therefore obtain significant fitness benefits from adjusting their brood sex ratio in response to the status of their pair bond. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

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.     

Offspring growth and mobility in response to variation in parental care: a comparison between populations

Life history theory emphasizes the importance of trade‐offs in how time and energy are allocated to the competing demands of growth, fecundity, and survival. However, avian studies have historically emphasized the importance of resource acquisition over resource allocation to explain geographic variation in fecundity, parental care, and offspring development. We compared the brood sizes and nestling mass and feather growth trajectories between orange‐crowned warblers Oreothlypis celata breeding in Alaska versus California, and used 24‐h video recordings to study the relationship between parental care and growth rates. Per‐offspring provisioning rates were highest in the smallest broods, and food delivery was positively correlated with nestling growth over the 24‐h period only in Alaska. Females in Alaska spent more time brooding, and juveniles there showed faster feather growth and earlier mobility compared with those in California. We also found differences in the energetic and nutritional content of insect larvae that could facilitate the observed differences in nestling growth relative to food provisioning. Our results point to the potential importance of food quality and parental provisioning of warmth, in addition to food, for explaining avian growth patterns. We highlight the need to quantify multiple dimensions of parental care and of offspring growth and development, and to better understand the relationships between feather growth, nestling period length, and fledgling mobility.     

Sex allocation and nestling survival in a dimorphic raptor: does size matter?

Fisher's theory predicts equal sex ratios at the end of parentalcare if the costs and benefits associated with raising eachsex of offspring are equal. In raptors, which display variousdegrees of reversed sexual size dimorphism (RSD; females thelarger sex), sex ratios biased in favor of smaller males areonly infrequently reported. This suggests that offspring ofeach sex may confer different fitness advantages to parents.We examined the relative returns associated with raising eachsex of offspring of the brown falcon Falco berigora, a medium-sizedfalcon exhibiting RSD (males approximately 75% of female bodymass) and subsequent sex ratios. Female nestlings hatched eitherfirst or second did not receive more food nor did they hatchfrom larger eggs or remain dependent on parents for longer periodsthan male offspring from these hatch orders. Together with previousstudies this result indicates that even in markedly dimorphicspecies, the required investment to raise the larger sex islikely to be less than that predicted by body size differencesalone. Moreover, among last-hatched nestlings, both sexes faceda reduced food allocation and suffered a slower growth rateand thus final body size, with a concurrent increased probabilityof mortality. For last-hatched females the reduction in foodallocation was more marked, with complete mortality of all last-hatchedfemale nestlings monitored in this study. Once independent,males of any size but only larger females are likely to be recruitedinto the breeding population. The sex-biased food allocationamong last-hatched offspring favoring males thus reflects therelative returns to parents in raising a small member of eachsex.     

Sexual pigmentation and parental risk‐taking in yellow warblers Setophaga petechia

Adult‐directed predation risk imposes important behavioral constraints on parents and might thus alter relationships between costly sexual ornaments and parental performance. For instance, under low predation risk, highly ornamented individuals might display better parental performance than others, as predicted by ‘good parent’ models of sexual selection. However, under high risk of predation, highly ornamented individuals might abandon parental effort if conspicuous to predators, or if social partners are more willing to take parental risks when paired with highly ornamented mates. We experimentally elevated perceived adult‐directed predation risk near nests to explore how carotenoid‐ and phaeomelanin‐based pigmentation in both sexes relate to parental risk‐taking for offspring in the yellow warbler Setophaga petechia. Compared to other males, males with more intense carotenoid‐based pigmentation maintained higher levels of paternal effort under predation risk at highly concealed nests, but reduced nestling provisioning rate more at exposed nests. Further, when faced with predation risk, females with more phaeomelanin‐based pigmentation reduced nestling provisioning rate less than other females, regardless of nest concealment. Females displayed higher parental effort across treatments when paired to males with more colorful carotenoid pigmentation. However, birds did not reduce parental effort under risk less when paired to a highly ornamented mate, suggesting that predation risk did not accentuate differential allocation. Males did not take fewer parental risks than females. Results indicate that nest concealment modifies parental risk‐taking by males with colorful carotenoid‐based pigmentation, and suggest that female melanin‐based pigmentation may indicate boldness and greater a propensity to take parental risks.     

Diet,maternal condition,and offspring sex ratio in the zebra finch,Poephila guttata

Where maternal condition affects condition and reproductive potential of offspring differentially with respect to sex, mothers in relatively good condition should produce more of the sex whose fitness is more dependent on condition. We experimentally manipulated body-condition in unmated zebra finches by feeding them for three months on high- or low-quality diets. Birds were then allowed to breed, while keeping the same diets. Females on the lower quality diet were in better condition and hatched significantly more males than females. Poorer condition females hatched an equal sex ratio. Chicks fed on the low-quality diet, but not on the high-quality diet, showed female-biased mortality. These results show that facultative sex ratio manipulation and sex-biased mortality can act together to produce extreme sex ratios in this vertebrate.     

Sex allocation in haplodiploids is mediated by egg size: evidence in the spider mite Tetranychus urticae Koch

Haplodiploid species display extraordinary sex ratios. However, a differential investment in male and female offspring might also be achieved by a differential provisioning of eggs, as observed in birds and lizards. We investigated this hypothesis in the haplodiploid spider mite Tetranychus urticae, which displays highly female-biased sex ratios. We show that egg size significantly determines not only larval size, juvenile survival and adult size, but also fertilization probability, as in marine invertebrates with external fertilization, so that female (fertilized) eggs are significantly larger than male (unfertilized) eggs. Moreover, females with on average larger eggs before fertilization produce a more female-biased sex ratio afterwards. Egg size thus mediates sex-specific egg provisioning, sex and offspring sex ratio. Finally, sex-specific egg provisioning has another major consequence: male eggs produced by mated mothers are smaller than male eggs produced by virgins, and this size difference persists in adults. Virgin females might thus have a (male) fitness advantage over mated females.     

The relationship between female brooding and male nestling provisioning: does climate underlie geographic variation in sex roles?

Comparative studies of populations occupying different environments can provide insights into the ecological conditions affecting differences in parental strategies, including the relative contributions of males and females. Male and female parental strategies reflect the interplay between ecological conditions, the contributions of the social mate, and the needs of offspring. Climate is expected to underlie geographic variation in incubation and brooding behavior, and can thereby affect both the absolute and relative contributions of each sex to other aspects of parental care such as offspring provisioning. However, geographic variation in brooding behavior has received much less attention than variation in incubation attentiveness or provisioning rates. We compared parental behavior during the nestling period in populations of orange‐crowned warblers Oreothlypis celata near the northern (64°N) and southern (33°N) boundaries of the breeding range. In Alaska, we found that males were responsible for the majority of food delivery whereas the sexes contributed equally to provisioning in California. Higher male provisioning in Alaska appeared to facilitate a higher proportion of time females spent brooding the nestlings. Surprisingly, differences in brooding between populations could not be explained by variation in ambient temperature, which was similar between populations during the nestling period. While these results represent a single population contrast, they suggest additional hypotheses for the ecological correlates and evolutionary drivers of geographic variation in brooding behavior, and the factors that shape the contributions of each sex.     

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.     

Experimentally-elevated testosterone, female parental care, and reproductive success in a songbird, the Dark-eyed Junco (Junco hyemalis)

In male dark-eyed juncos (Junco hyemalis), experimentally elevated testosterone (T) decreases male parental care and offspring survival, but results in higher overall fitness through greater mating success. To help address the ensuing question of what prevents selection from favoring higher levels of T in male juncos, we manipulated T in female juncos. A previous study demonstrated no effect of experimentally elevated T on female incubation behavior, suggesting that female parental behavior might be insensitive to T. In this study we asked whether experimentally elevated T mediates other female parental behaviors and whether variation in T-mediated parental behavior might influence reproductive success. We videotaped free-living control- and T-females during nesting to quantify brooding behavior when young were 3 days old and provisioning behavior when young were 6 days old. Nest defense was measured by quantifying responses to a mounted predator placed near the nest. Reproductive success was assessed via fecundity, nestling quality, and nest survival. T-females spent less time than control females brooding but did not differ in provisioning rate. T-females performed fewer dives at the predator mount and, unlike controls, failed to increase defense as nesting progressed. T-females also had lower daily nest survival and lower nest success (odds of producing at least one fledgling). We conclude that some aspects of female parental behavior are sensitive to experimentally elevated T while others are not and consider the implications for the evolution of T-mediated characters in both sexes.     

The discovery of hatching and transition to feeding young by male Mountain Bluebirds

In many bird species, only females incubate the eggs, but both sexes feed nestlings. The means by which males of such species discover hatching and transition to feeding their offspring remains almost completely unexplored. Of particular interest are species with nests whose contents are concealed from view. During June and early July 2015 in the Bighorn Mountains of Wyoming, we used continuous video‐recording of nests of cavity‐nesting Mountain Bluebirds (Sialia currucoides) to document the transition to feeding young by males. We saw no evidence that females used distinct vocal or visual displays to signal hatching to males. Observing mates carrying eggshells away from, or food into, nest boxes did not appear to trigger provisioning by males. Rather, at all 24 nests observed, males did not begin feeding until they had come to nest boxes and presumably sensed the presence of hatchings directly. Individual males varied, however, in both the manner in which they inspected nest contents and the number of times they did so before starting to feed young. Although most males fully entered nest boxes where they could see, touch, hear, and possibly smell hatchlings (or eggshell parts), other males may have detected hatchlings only by sound or possibly smell while perched at a nest‐box entrance. Based on past studies of mice and doves, we suggest that, for provisioning behavior to begin, some kind of direct sensation of offspring may be necessary to activate relevant neurons in the medial preoptic area of the hypothalamus of males, an area of the brain important in parental care. Additional research is necessary to test this, and to examine the effects of factors such as hormone levels and breeding experience on the means and rapidity by which males discover hatching and transition to nestling provisioning.     

Complex sex allocation in the laughing kookaburra

In groups of the cooperatively breeding laughing kookaburra(Dacelo novaeguineae), offspring sex varied with the type ofsocial group and with hatch rank. Groups with female helpers,especially if all helpers were female, had male-biased clutchand fledging sex ratios. Groups without female helpers (unassistedpairs or male-only helpers) had female-biased clutch and fledgingsex ratios. Breeding females responded facultatively to increasesin the number of female helpers in their group by producingmore male eggs. These biases may occur if breeding femalestry to limit the number of daughters recruited into their groupbecause unlike male helpers, female helpers depress the breedingsuccess of their parents. Across all nests, two-thirds of first-hatchedyoung were male, two-thirds of second-hatched young were female, and the sex ratio of third-hatched young was even. Hatch ranksex ratios also varied dramatically between different typesof social groups, from 16.7% for second-hatched nestlings ofunassisted pairs to 100% for first-hatched nestlings of groupswith only female helpers. A corollary of the relationship betweenhatch rank and sex was that hatching sex sequences were distributed nonrandomly: all groups avoided hatching a daughter first followedby a son (FM). Sibling competition is aggressive and sometimesfatal. Since females grow to be 15% larger than males the hatchingsequence of sexes could affect nestling growth and mortality.However, an exhaustive analysis found little evidence thatgrowth or survival of males was compromised if hatched aftera sister. The small number of FM sequences may only have occurredin nests that were able to ameliorate any negative consequences.Alternatively, when clutch size is small and fledging successunpredictable because of brood reduction, the preferred broodsex ratio may be contingent on the number of fledged young,making it advantageous to order the sexes in the brood.     

The post‐fledging period in a tropical bird: patterns of parental care and survival

How environmental conditions affect the timing and extent of parental care is a fundamental question in comparative studies of life histories. The post‐fledging period is deemed critical for offspring fitness, yet few studies have examined this period, particularly in tropical birds. Tropical birds are predicted to have extended parental care during the post‐fledging period and this period may be key to understanding geographic variation in avian reproductive strategies. We studied a neotropical passerine, the western slaty‐antshrike Thamnophilus atrinucha, and predicted greater care and higher survival during the post‐fledging period compared to earlier stages. Furthermore, we predicted that duration of post‐fledging parental care and survival would be at the upper end of the distribution for Northern Hemisphere passerines. Correspondingly, we observed that provisioning continued for 6–12 weeks after fledging. In addition, provisioning rate was greater after fledging and offspring survival from fledging to independence was 75%, greater than all estimates from north‐temperate passerines. Intervals between nesting attempts were longer when the first brood produced successful fledglings compared to nests where offspring died either in the nest or upon fledging. Parents delayed initiating second nests after the first successful brood until fledglings were near independence. Our results indicate that parents provide greater care after fledging and this extended care likely increased offspring survival. Moreover, our findings of extended post‐fledging parental care and higher post‐fledging survival compared to Northern Hemisphere species have implications for understanding latitudinal variation in reproductive effort and parental investment strategies.