Vocal begging by nestlings and vulnerability to nest predation in Meadow Pipits Anthus pratensis; to what extent do predation costs of begging exist?

Models of the evolution of parent-offspring communication and of brood size assume that the noisy begging by the young is costly in terms of predation. The present study was designed to investigate cause-and-effect relations between the behaviour of the offspring and adult Meadow Pipits Anthus pratensis and the risk of nest predation. Harriers ( Circus spp.), which can use auditory signals, were the main predators in the study area. I found that the intensity of vocal begging by broods did not influence their survival, despite the fact that the proportion of time that parents sacrificed to nest guarding declined with increased begging and some broods readily begged if a stimulus resembling the approaching parent was produced. The lack of fitness penalties for noisy broods was attributed to the following factors: (1) the open habitat was easy to scan, and parents silenced the young before feeding them if a predator was close (this adaptation was demonstrated in an experiment) and (2) parent birds could not deter harriers searching for prey, and therefore nest guarding did not influence the probability of nest detection. Pitfalls in the interpretation of relations between the vocal begging and the vulnerability to nest predation are discussed.     

Offspring dynamics affect food provisioning,growth and mortality in a brood-caring spider

In brood-caring species, family members are faced with a conflict over resource distribution. While parents are selected to adapt the amount of care according to their offspring''s needs, offspring might be selected to demand more care than optimal for parents. Recent studies on birds have shown that the social network structure of offspring affects the amount of care and thus the fitness of families. Such a network structure of repeated interactions is probably influenced by within-brood relatedness. We experimentally manipulated the group composition in a brood-caring spider to test how the presence of unrelated spiderlings affects the dynamics between female and brood as well as within broods. Broods consisting of siblings grew better and had a lower mortality compared with mixed broods, no matter whether the caring female was a genetic or foster mother. Interestingly, we found that foster mothers lost weight when caring for sibling broods, whereas females caring for mixed broods gained weight. This indicates that females may be willing to share more prey when the brood contains exclusively siblings even if the entire brood is unrelated to the female. Resource distribution may thus be negotiated by offspring dynamics that could have a signalling function to females.     

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.     

Do honest signalling models of offspring solicitation apply to insects?

Honest signalling models predict that the intensity of solicitation by offspring influences the level of provisioning provided by parents and reflects offspring need. The empirical evidence supporting these predictions primarily comes from studies of birds or mammals. Thus, although parental care of altricial offspring is taxonomically widespread, the generality of these models is not well known. To investigate whether honest signalling models apply to insects, we manipulated parent and offspring behaviour in the burying beetle Nicrophorus orbicollis, a species with advanced parental care. First, within biparental care, we manipulated the brood size to alter the parents'' perception of offspring need. We measured the care giving behaviour of male and female parents to examine whether either adjusts its level of care according to offspring need. In the second experiment, because two parents together provision the brood more often than single parents, we manipulated the number of care givers (uniparental and biparental care) and measured offspring solicitation to assess whether offspring change their behaviour in response to need. Our results show that parent behaviour is broadly consistent with the first prediction of the models; both sexes provisioned larger broods more often than smaller broods. Larval solicitation was also consistent with the second prediction; larvae that were provisioned less often begged more. Our results provide evidence that honest signalling models can be applied to insects as well as vertebrates, although there are also subtle differences in care giving behaviour that may be important.     

EVOLUTION OF OBLIGATE SIBLICIDE IN BOOBIES. 2: FOOD LIMITATION AND PARENT–OFFSPRING CONFLICT

Proximate limitation on parental food delivery has long been invoked to explain the evolution of single-chick broods of pelagic seabirds such as masked boobies (Sula dactylatra). A second possible proximate limit on brood size is siblicide driven by genetic parent–offspring conflict (POC) over brood size, if siblicidal offspring can reduce brood size to one even if the parents' optimal brood size is greater than one. I tested these two hypotheses by experimentally suppressing obligate siblicide in masked booby broods and comparing breeding parameters of these broods with unmanipulated single-chick control broods. Per capita mortality rate of experimental nestlings was higher than that of controls, but this deficit was more than made up by larger brood size. Parents of experimental broods brought more food to offspring, had higher fledging success, and apparently incurred no additional major short-term cost of reproduction, relative to parents of control broods, thus refuting the food limitation hypothesis. Estimates of inclusive fitness of chicks in experimental broods were higher than were those of control nestlings, a result inconsistent with the POC hypothesis that the siblicidal offspring's optimal brood size is one while the parents' optimum is greater than one. This discrepency between natural brood size and apparent brood size optima might be resolved in several ways: experimental artifacts may give misleading estimates of optimal brood size; experimental and control offspring may have different reproductive values at the time of fledging; nestling masked boobies may face a special frequency-dependent case of POC in which the high risk of sharing a nest with a siblicidal sibling makes invasion of other behavioral genotypes difficult even when offspring and parent inclusive fitnesses are higher from a nonsiblicidal brood of two than from a brood of one.     

Does resource availability affect offspring begging and parental provisioning in a partially begging species?

In species where parents repeatedly provide their offspring with food, the offspring often communicate their need to the parents. Burying beetles, which breed on a wide size range of carcasses of small vertebrates, are interesting model systems to test theories on begging, because the larvae show partial begging, that is, they obtain food through both signalling to their parents (begging) and feeding directly from the carcass. We manipulated resource availability inNicrophorus vespilloides by providing parents with mouse carcasses spanning a wide size range, and allowing them to rear the larvae that hatched, so that both the amount of resources and the number of siblings varied. Time spent begging by each larva was strongly influenced by the time parents spent near the larvae. Brood size had a nonlinear effect on larval begging, with begging increasing with brood size for relatively smaller broods and decreasing again for larger broods. Carcass size and number of parents present had no effect on begging. Time spent provisioning the larvae by the parents was strongly associated with the time spent begging by each larva. Parents spent more time provisioning under biparental care than under uniparental care, while brood size and carcass size had no significant effect. These findings suggest that the larvae adjust their begging to the behaviour of their parents and the number of siblings, but not to the amount of resources. Furthermore, parents adjust the time spent provisioning to the average amount of begging by each larva in the brood, and not to the availability of resources.     

Sense and sensitivity: responsiveness to offspring signals varies with the parents' potential to breed again

How sensitive should parents be to the demands of their young? Offspring are under selection to seek more investment than is optimal for parents to supply, which makes parents vulnerable to losing future fitness by responding to manipulative displays. Yet, parents cannot afford to ignore begging and risk allocating resources inefficiently. Here, we show that parents may solve this problem by adjusting their sensitivity to begging behaviour in relation to their own likelihood of breeding again, a factor largely neglected in previous analyses of parent–offspring interactions. In two carotenoid-supplementation experiments on a New Zealand passerine, the hihi Notiomystis cincta, we supplemented adults to enhance their propensity to breed again, and supplemented entire broods to increase their mouth colour, thus enhancing their solicitation display. We found that adults that attempted two breeding attempts a season were largely insensitive to the experimentally carotenoid-rich gapes of their brood, whereas those that bred just once responded by increasing their rate of provisioning at the nest. Our results show that parents can strategically vary their sensitivity to begging in relation to their future reproductive potential. By restricting opportunities for offspring to influence provisioning decisions, parents greatly limit the potential for offspring to win parent–offspring conflict.     

Cooperation, conflict, and crèching behavior in goldeneye ducks

Crèching behavior, or brood amalgamation, results in offspring being reared by adults other than their genetic parents. Although a variety of hypotheses have been proposed to explain this behavior, most assume either that brood amalgamation is accidental (i.e., nonselected) or that adoption of young is selected for because of social benefits to the young and/or adopting parents. We propose, instead, that brood amalgamation is a function of two separate processes: brood desertion and brood adoption. To examine brood desertion, we develop a graphic model to predict when parents should abandon their young and we test this model experimentally for the Barrow's goldeneye (Bucephala islandica). As predicted, females deserted their offspring when the size of the brood was experimentally reduced. Brood adoption occurred when deserted ducklings joined other broods. However, the success of ducklings in doing so was strongly dependent on the availability of potential host broods and on the age of the recipient broods. Foreign ducklings were readily accepted into young broods (<10 d old) but invariably were rejected from old broods. We could detect no benefits or costs of brood adoption to the host females, contrary to the expectations of a social benefit hypothesis. Our experiments indicate that Crèching behavior is driven by selection on adults to abandon their brood when the benefits of continued investment are outweighed by the reduction in future reproduction and selection on deserted ducklings to join other broods to obtain parental care. Rather than a form of cooperative brood care, Crèching in goldeneyes is perhaps best considered as a form of reproductive parasitism, entailing parent-offspring conflict over brood desertion and intergenerational conflict over adoption of abandoned young.     

Calling at a cost: elevated nestling calling attracts predators to active nests

Begging by nestling birds has been used to test evolutionary models of signalling but theory has outstripped evidence. Eavesdropping predators potentially impose a cost on begging that ensures signal honesty, yet little experimental evidence exists for such a cost at active nests because the use of artificial nests, long playback bouts and absence of parents may have exaggerated costs. We broadcast short periods (1 h) of either nestling vocalizations or background noise at active white-browed scrubwren, Sericornis frontalis, nests. Nestlings called naturally during both treatments, allowing us to test whether elevated calling increases risk, a key but rarely tested assumption of evolutionary models. Predators visited nests exclusively during periods of elevated calling. Furthermore, playbacks affected neither adult visits nor nestling activity, suggesting that calling alone attracted predators. Adults gave alarm calls and nestlings usually called less when predators approached nests. Predation risk to broods is, therefore, likely to fluctuate substantially over short periods of time, depending on nestling hunger and whether adults or young have detected predators. This study confirms a present-day cost of nestling begging, demonstrates that this cost can be incurred over short periods and supports the importance of parent-offspring antipredator strategies in reducing predation risk.     

Ectoparasites and reproductive trade-offs in the barn swallow (Hirundo rustica)

Parents are predicted to trade offspring number and quality against the costs of reproduction. In altricial birds, parasites can mediate these costs because intensity of parasitism may increase with parental effort. In addition, parasites may mediate a trade-off between offspring number and quality because nestlings in large broods may have reduced anti-parasite immune defence. In this study, we experimentally analysed the effect of brood size on infestation by an ectoparasitic mite in nests of barn swallows (Hirundo rustica). Nests with an enlarged brood had larger prevalence and intensity of infestation than those with a reduced brood. Importantly, each nestling in enlarged broods was exposed to a larger number of mites, even when measured on a per nestling basis, than in reduced broods. Nestlings in enlarged broods had smaller body mass and T-cell-mediated immune response compared to reduced broods. T-cell-mediated immune response and feather growth were negatively correlated with per nestling intensity of infestation in enlarged but not in reduced broods. The results suggest that nestlings in enlarged broods have depressed immunity leading to larger per nestling mite infestation. Hence, exposure to parasites of offspring and parents increases with brood size, and parasitism can thus mediate trade-offs between reproduction and number and quality of the progeny in the barn swallow.     

Parental allocation of food to nestling tree swallows: the influence of nestling behaviour, sex and paternity

Parents are expected to invest more in young that provide the greatest fitness returns. The cues that parents use to allocate resources between their offspring have received much recent attention. In birds, parents may use begging intensity, position in the nest or nestling size as cues to provision the most competitive young or those most likely to survive. It may also benefit parents to invest in young differentially by sex or relatedness if the fitness returns of sons and daughters differ or broods are sired by multiple males. We examined the allocation of food to tree swallow, Tachycineta bicolor, nestlings in relation to their begging behaviour, size, sex and paternity. Provisioning by parents was not related to nestling size, sex or paternity. The begging behaviour of nestlings did not differ with respect to sex or paternity. Both parents were more likely to feed nestlings that begged first or were closer to the nest entrance, suggesting that parents allocate food resources in response to cues that nestlings control. As a consequence, brood reduction was facilitated by biased provisioning within the brood in addition to the nestling size hierarchies created by hatching asynchrony. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Interaction between parental care and sibling competition: parents enhance offspring growth and exacerbate sibling competition

Species with elaborate parental care often also show intense sibling competition over resources provided by parents, suggesting joint evolution of these two traits. Despite this, the evolution of elaborate parental care and the evolution of intense sibling competition are often studied separately. Here, we examine the interaction between parental food provisioning and sibling competition for resources through the joint manipulation of the presence or absence of parents and brood size in a species with facultative parental care: the burying beetle Nicrophorus vespilloides. The effect of the interaction between the presence or absence of parents and brood size was strong; brood size had a strong effect on growth when parents provided care, but no effect when parents were absent. As in previous studies, offspring grew faster when parents were present than when parents were absent, and offspring grew faster in smaller broods than in larger broods. Our behavioral observations showed that brood size had a negative effect on both the amount of time parents spent providing resources to individual offspring and the offspring's effectiveness of begging, confirming that the level of sibling competition increased with brood size. Furthermore, offspring in larger broods shifted more from begging toward self-feeding as they grew older compared to offspring in small broods. Our study provides novel insights into the joint evolution of parental care and sibling competition, and the evolution of offspring begging signals. We discuss the implications of our results in light of recent theoretical work on the evolution of parental care, sibling competition, and offspring begging signals.     

Cichlid-catfish mutualistic defense of young in Lake Malawi,Africa

Summary Bagrid catfish and cichlid parents engage in a mutualistic defense of their young from predators. Over 50% of the catfish broods observed contained cichlid young, primarily of three species: Crytocara pleurostigmoides, C. pictus and Rhamphocromis sp. Three catfish broods, monitored for over 50 days, had a survivorship rate 6 times greater during periods when cichlids were present than when cichlids were absent. During two and a half h of observations of catfish broods without cichlids, I observed 23 strikes by predators on catfish young. However, when cichlids were present no foraging attacks were observed upon the catfish, but 32 occurred against cichlids in two and a half hours of observations. When the catfish parents were experimently removed the cichlid young were consumed first then the catfish young. When cichlids were present the catfish young survived over 80% longer than those in broods without cichlids. It is concluded that the parental catfish increase the survival of their own young be allowing cichlids into the brood. Because of an asymmetry of costs between the two species, cichlid mothers are more likely to abandon these interspecific broods than are the catfish.     

Premating Behavior in the Subgenus Limia (Pisces: Poeciliidae): Sexual Selection and the Evolution of Courtship

A three-year field-study of Richardson's ground squirrels was conducted to assess whether alarm calling functions to warn close relatives (“kin selection” hypothesis) or manipulate conspecifics (a “selfish” hypothesis). S. richardsonii had distinct calls for terrestrial and aerial predators, and the responses of squirrels varied appropriately according to the context of calls, implying that calling conveyed correct information concerning the nature of the danger. Alarm calling elicited by naturally occurring encounters with potential predators during 454 h of observation, and by a thrown frisbee in 70 experimental trials, was not equally probable for all age/sex classes. Squirrels were most likely to call when they had offspring or siblings nearby, which is supportive of the hypothesis that alarm calling is maintained by kin selection. Adult males, residing in the vicinity of either their probable progeny or their nonlittermate half-siblings, were the most likely age/sex class to call during the lactation period when young were below ground and were most vulnerable. I conclude that alarm calling by Richardson's ground squirrels is nepotistic rather than manipulative.     

Brood amalgamation in the Bristle-thighed Curlew Numenius tahitiensis: process and function

Alloparental care in birds generally involves nonbreeding adults that help at nests or breeding adults that help raise young in communal nests. A less often reported form involves the amalgamation of broods, where one or more adults care for young that are not their own. We observed this phenomenon among Bristle-thighed Curlew Numenius tahitiensis broods in western Alaska during 1990–1992. Amalgamation of broods generally involved the formation of temporary and extended associations. Temporary associations were formed by the incidental convergence of broods soon after they left their nests. During this period, parents defended distinct brood-rearing areas, were antagonistic to conspecifics and remained together for less than 3 days. Extended associations formed when chicks were 1–2 weeks old. Here, parents and their broods occupied distinct, but adjacent, brood-rearing areas and moved around as a unit. Whether a brood participated in either temporary or extended associations or remained solitary appeared to depend on brood density in the immediate area and on hatching date. When chicks were 3–4 weeks old, aggregations of up to ten broods formed wherein young mixed and parents defended a common brood-rearing area. All broods (n = 48) that survived to fledging joined such aggregations. Alloparental care involved only antipredator defence and was not associated with activities such as feeding and brooding. Most female parents abandoned their broods shortly after the young could fly and when aggregations were forming. The female parent of a pair always deserted its young before or on the same day as the male parent and, in every aggregation, one or two males continued to tend young for about 5 days longer than other male parents. In most cases, adults deserted the young 2–6 days before the young departed the area when about 38 days old. Bristle-thighed Curlews also formed temporary associations with American and Pacific Golden Plover Pluvialis dominica and Pluvialis fulva, Whimbrel Numenius phaeopus, Bar-tailed Godwit Limosa lapponica, Western Sandpiper Cal-idris mauri and Long-tailed Skua Stercorarius longicaudus. Curlews and other larger bodied species commonly attack-mobbed predators together, whereas smaller bodied species generally gave alarm calls and circled the predators. For all species, the intensity of antipredator defence by attending adults gradually decreased as young became older and aggregations formed. We suggest that amalgamation of broods among Bristle-thighed Curlew enhances predator defence, aids in the process of flock formation for migrating young, and allows females and some males to desert their young earlier.     

Interspecific Interactions and the Scope for Parent-Offspring Conflict: High Mite Density Temporarily Changes the Trade-Off between Offspring Size and Number in the Burying Beetle,Nicrophorus vespilloides

Parents have a limited amount of resources to invest in reproduction and commonly trade-off how much they invest in offspring size (or quality) versus brood size. A negative relationship between offspring size and number has been shown in numerous taxa and it underpins evolutionary conflicts of interest between parents and their young. For example, previous work on vertebrates shows that selection favours mothers that produce more offspring, at the expense of individual offspring size, yet favours offspring that have relatively few siblings and therefore attain a greater size at independence. Here we analyse how this trade-off is temporarily affected by stochastic variation in the intensity of interspecific interactions. We examined the effect of the mite Poecilochirus carabi on the relationship between offspring size and number in the burying beetle, Nicrophorus vespilloides. We manipulated the initial number of mites in the reproductive event (by introducing either no mites, 4 mites, 10 mites, or 16 mites), and assessed the effect on the brood. We found a similar trade-off between offspring size and number in all treatments, except in the ''16 mite'' treatment where the correlation between offspring number and size flattened considerably. This effect arose because larvae in small broods failed to attain a high mass by dispersal. Our results show that variation in the intensity of interspecific interactions can temporarily change the strength of the trade-off between offspring size and number. In this study, high densities of mites prevented individual offspring from attaining their optimal weight, thus potentially temporarily biasing the outcome of parent-offspring conflict in favour of parents.     

Alarm calls of the Cyprus Wheatear Oenanthe cypriaca—one for nest defence, one for parent–offspring communication?

Alarm calling has been interpreted from the viewpoint of parental investment and nest defence. Calling should increase with an increasing value of the brood to the caller or with increasing vulnerability of the brood. Parental alarm calling might also be viewed from assessment–management with callers using vocalisations to elicit affective responses in others, e.g. in silencing offspring. I examined parental alarm calling in the Cyprus Wheatear (Oenanthe cypriaca) at different developmental stages. The Cyprus Wheatear produces two alarm calls, but one is emitted only during a short period of the year (type II alarm call). The commoner alarm call (type I alarm call), however, is used year-round. Predation was simulated using consistent approach by human observers. The brood cycle was divided into (I) incubation, (II) nestling, (III) early fledging, and (IV) late fledging. Type I calls increased from phases I to III, and decreased afterwards. Type II calls were absent during incubating and occurred during nestling and early fledging stages. As a conclusion, I suggest that the type I alarm call might be used to alert the mate, deter the predator and signal strength of nest defence, while the type II alarm call is addressed to offspring.     

Experimental study of alarm calls of the oriental tit (Parus minor) toward different predators and reactions they induce in nestlings

Anti-predatory strategies of birds are diverse and may include predator-specific alarm calls. For example, oriental tit (Parus minor) parents can distinguish snakes from other predators and produce snake-specific referential vocalizations ("jar" call) when a snake poses a threat to their nest. The “jar” call has a very specific function to induce fledging of nestlings close to fledging age. This reaction ensures nestlings' survival in natural encounters with snakes that are capable of entering nest cavities and kill entire broods. Sciurid rodents, like chipmunks, may pose a similar threat to cavity-nesting birds. We explored the hypothesis that parents use the fledging-inducing alarm vocalizations in this situation, because chipmunks, like snakes, can kill the brood upon entering the nest cavity. We compared alarm calls of parents toward two predators (chipmunk and snake) who pose a similar threat to the nestlings in a nest cavity, and toward an avian predator (Eurasian jay) who cannot enter nest cavities and poses no threat to the nestlings in a nest. Our results show that the vocal responses of oriental tits were different among the three predators. This suggests that the acoustic properties of vocal responses to predators are different between predators of a similar hunting strategy (nest-cavity entering). The playback of recorded vocal responses of parents to chipmunks did not trigger the fledging of old nestlings, whereas the vocalizations toward a snake did, as shown by earlier studies. Our study suggests that the vocal response of parents does not carry information about the ability of predators to enter the nest cavity and confirms the special status of alarm calls triggered by snakes.     

Azure‐winged Magpies Cyanopica cyanus trade off reproductive success and parental care by establishing a size hierarchy among nestlings

It is common in birds that the sizes of nestlings vary greatly when multiple young are produced in one nest. However, the methods used by parents to establish size hierarchy among nestlings and their effect on parental provisioning pattern may differ between species. In the Azure‐winged Magpie Cyanopica cyanus, we explored how and why parents controlled the sizes of nestlings. Asynchronous hatching was the main cause of size hierarchy within the brood, although the laying of larger eggs later in the laying sequence reduced this effect. Parents with asynchronous broods produced more eggs and fledged more nestlings than those with synchronous broods but their brood provisioning rates, food delivery per feeding bout and feeding efficiency did not differ. We performed a cross‐fostering experiment to synchronize some asynchronous broods. Provisioning rates of asynchronous broods were lower than those of synchronized broods, but the daily growth rates and fledging body mass of their nestlings were not different. Our findings indicate that parents of asynchronous broods can achieve higher reproductive success than those of synchronous broods based on the same parental care, and the same reproductive success as those of synchronized broods based on less parental care. It appears that parent birds can better trade off reproductive success and parental care by establishing a size hierarchy among nestlings.     

Paternal influence on growth and survival of dark-eyed junco young: do parental males benefit?

Recent studies of typically monogamous passerine birds have suggested that the fitness benefits males derive by caring for their young may not be as great as was previously thought. This study was conducted to determine whether parental care by male dark-eyed juncos, Junco hyemalis, serves to increase either the quantity or quality of young that they produce. Over a 4-year period, males were caught at the time their eggs hatched, and the subsequent growth and survival of the young of unaided females and control pairs were compared. Broods raised by unaided females gained body mass more slowly and fledged at slightly lower mass than those raised by two parents. However, fledging mass was not correlated with survival to independence. There were no differences in tarsus growth between the two treatment groups. Entire brood loss to predators occurred as often among females without male help as it did among those with male help. However, partial brood loss was more common among female-only broods than among controls; this difference was largely attributable to higher rates of starvation and exposure in female-only broods. There appeared to be an interaction between growth and predation. Female-only broods that were below the median mass of combined treatment groups at 5 days of age were more likely than all other broods to experience partial or complete predation. Male impact on offspring survival varied with age of the offspring. When years were combined, males tended to increase survival during the first half of the nestling period, but their impact at the time of nest-leaving was minimal. In all years, from nest-leaving to independence (ca. 2 weeks), broods without male help survived only about half as well as did those with male help. Independent young raised by one parent were as likely to return the following spring as were young raised by two parents. Thus, paternal care benefits males by improving the survivorship of their fledglings, and may also act as a buffer against poor female parental quality and inclement weather. However, the magnitude of these benefits is such that bigamous males might achieve greater reproductive success than monogamous males. Various possible male strategies are discussed.