Need and nestmates affect begging in tree swallows

We conducted an experiment on nestling tree swallows (Tachycineta bicolor) to examine predictions from signalling models for the evolution of conspicuous begging behaviour. Specifically, we examined the relationship between (1) nestling begging intensity and hunger, (2) begging intensity and parental provisioning and (3) begging intensity and nestmate condition. Forty broods of 9-day-old nestlings were removed from their nests for 1 h and assigned to one of the following three treatments: (1) all nestlings in the brood deprived of food (n = 13), (2) all nestlings in the brood fed (n = 11) or (3) half the nestlings in the brood deprived and half fed (n = 16). Videotapes before and after the treatments showed that begging intensity increased in broods in which all of the nestlings had been deprived and decreased in broods in which all of the nestlings had been fed. Deprived nestlings in the half-and-half treatment did not change their begging intensity in response to treatment, while fed nestlings in this treatment group showed a decrease in begging intensity. Parent tree swallows increased their feeding rate to deprived broods and decreased their rate to fed broods. Within broods, parents decreased their feeding rate to fed nestlings, but showed no significant change in feeding to deprived nestlings. Our results suggest that begging intensity is influenced by hunger and that parents appear to respond to variation in begging intensity. The begging of nestmates also appears to influence begging independently of need. These results are consistent with predictions derived from signalling models of begging. Received: 20 June 1997 / Accepted after revision: 19 January 1998     

Begging calls and parental feeding decisions in tree swallows (Tachycineta bicolor)

We conducted playback experiments to examine how parent tree swallows (Tachycineta bicolor) use nestling begging calls to distribute feedings to individuals within broods. In a first study, we used a paired-choice test to determine if parents discriminated between the taped begging calls of nestlings deprived of food and those of nestlings that had been recently fed. Our results showed that parents directed their first feeding attempt towards model nestlings near speakers playing deprived calls significantly more often than to models near speakers playing fed calls. They also made more feeding attempts overall to models with deprived calls. In the second study, we varied call rate and amplitude to examine which call features parents might use to discriminate begging calls. Parents directed significantly more first feeding attempts and more feeding attempts overall towards non-begging nestlings near speakers playing high call rates than to nestlings near speakers playing low call rates. They did not, however, discriminate between calls differing in amplitude. Previous studies have shown that parent birds use begging calls to regulate overall feeding rates to the brood. Our results suggest that parent tree swallows also use begging calls when feeding individual nestlings and, in particular, prefer calls associated with increased levels of nestling hunger. Received: 14 February 2000 / Revised: 6 October 2000 / Accepted: 16 October 2000     

Provisioning rules in tree swallows

Conflict between parents and offspring may result in offspring exaggerating their needs and parents devaluing their begging signals. To determine whether this occurs, it is first necessary to establish the link between need, begging and parental response. The purpose of our study was to examine these relationships in tree swallows (Tachycineta bicolor). Parents preferentially fed nestlings that begged sooner, reached higher and were closer to the front of the nestbox (Fig. 1). Begging intensity of both individuals and entire broods increased with relatively long periods between feeding visits. Within broods, parents responded to increased begging intensity by increasing their feeding rate, although this effect was relatively weak. Large and small nestlings did not differ in their begging behavior and all nestlings, regardless of size, were fed at similar rates. Despite the overall equity in feeding, male parents preferentially fed larger nestlings while female parents fed smaller nestlings. Nestlings did not increase their begging intensity in response to begging by nestmates. Our results suggest that begging is related to need in this species and that parents respond to variation in begging intensity. Received: 4 May 1995/Accepted after revision: 17 December 1995     

Nestling American robins compete with siblings by begging

Summary The evolution of intense begging by dependent nestling birds has recently been the subject of several theoretical papers. The interesting problem here is that nestlings should be able to communicate their nutritional status to parents in ways that are less costly energetically and less likely to attract predators. Thus, conspicuous begging behaviour is thought to have evolved as a result of either competition among nestmates or the manipulation of their parents to provide more food than would otherwise be favoured by selection. We studied sibling competition for parental feedings in the American robin (Turdus migratorius). We demonstrate that the probability that an individual nestling received food was related to several indices of begging. When we experimentally prevented parents from feeding part of their brood, both the intensity of begging and the number of feedings subsequently received by food-deprived nestlings increased. Furthermore, the begging intensity of those nestlings that were not food-deprived also increased in response to the begging of their hungrier siblings.Offprint requests to: R. Montgomerie     

Begging signals more than just short-term need: cryptic effects of brood size in the pied flycatcher (Ficedula hypoleuca)

The begging of nestling birds is known to reliably signal short-term nutritional need, which is used by parents to adjust rates of food delivery and patterns of food distribution within broods. To test whether begging signals reflect more than just short-term feeding history, we experimentally created 18 "small" (4-nestling) and 18 "large" (8-nestling) broods in the pied flycatcher (Ficedula hypoleuca). Compared to small broods, large broods were provisioned by parents at a greater rate, but at a lower visit rate per nestling and with no obvious differences in load mass per visit. However, lower rates of food mass delivery per nestling in large broods did not result in any measurable reduction in nestling growth (i.e. "long-term need") or in any increase in the begging effort per individual nestling whilst in the nest. Mid-way through the nestling period we also used hand-feeding laboratory trials to assess in more detail individual begging behaviour and digestive performance of the three mid-ranking nestlings from each brood. More food items were required at the start of each trial to satiate nestlings from large broods, but despite this initial control for "short-term need", nestlings from large broods went on to beg at consistently higher rates and at different acoustic frequencies. Large brood nestlings also produced smaller faecal sacs, which were quantitatively different in content but did not differ in frequency. We suggest that different nutritional histories can produce cryptic changes in nestling digestive function, and that these can lead to important differences in begging signals despite controlling for short term need.     

Parent-absent signalling of need and its consequences for sibling competition in the barn swallow

In species with parental care, competition among siblings for access to limited parental resources is common. Sibling competition can be mediated by begging behaviour, a suite of different visual and acoustic displays by which offspring solicit parental care. These are mostly addressed to the parents upon food provisioning, but can also be performed in the absence of the attending parents. This so-called parent-absent begging (PAB) may function as an intrabrood communication signal and potentially affect intrabrood competition dynamics for access to food. We investigated the role of PAB in moulding sibling interactions and its effect on food intake among altricial barn swallow (Hirundo rustica) nestlings, both under normal and experimentally reduced food intake. Frequency of PAB increased after food deprivation. Nestlings that had performed PAB increased their begging intensity upon the subsequent parental feeding visit, while siblings reduced their own begging level, but only when they had not been food-deprived. As a consequence, nestlings which had performed PAB before parental arrival had larger chances of receiving food. However, nestlings did not benefit from displaying PAB when competing with food-deprived siblings. Our findings show that PAB reliably reflects need of food, indicating that a nestling will vigorously compete for the subsequent food item. By eavesdropping siblings' PAB displays, nestlings may optimally balance the costs of scrambling competition, the direct fitness gains of being fed and the indirect fitness costs of subtracting food to needy kin. However, large asymmetries in satiation between competitors may lead individual offspring to monopolize parental resources, irrespective of PAB displays.     

Dynamics of calling by tree swallow (Tachycineta bicolor) nestmates

The begging display of nestling passerine birds has become a model for examining the evolution of animal signals. A particular problem for nestlings when transmitting begging signals to parents may be interference from nestmates. The strategies used by nestlings to reduce signal interference have not been studied, yet potentially contribute to the design of these complex displays. In this study, we recorded the begging calls of nestling tree swallows (Tachycineta bicolor) when alone and with a begging nestmate, to determine whether nestlings changed the output, structure or timing of their calls in ways that would reduce acoustic interference. We found that nestlings increased their call rate in the presence of a begging nestmate, but did not alter the length, amplitude or frequency of their calls. They also appeared not to adjust the timing of their calls to avoid those of nestmates. Contrary to expectation, nestling calls became more similar in some aspects when nestmates called together. An increase in call rate in the presence of a begging nestmate should increase the likelihood that a parent detects an individual's calls. However, if all nestlings increase their calling rate in response to competitors, then the overall level of acoustic interference across the brood is potentially increased, an effect exacerbated by the tendency for call similarity to increase when calling together. We discuss how increasing call rate may improve detectability despite this effect and we also examine how an increase in rate and call similarity may serve to produce a strong brood signal.     

The effect of pup vocalisations on food allocation in a cooperative mammal, the meerkat (Suricata suricatta)

In the meerkat (Suricata suricatta), a cooperative mongoose, pups follow potential feeders while the group is foraging and emit incessant calls when soliciting food from them. In contrast to a ’stationary’ brood of chicks, in which nestlings are fed at a fixed location, meerkat pups are ’mobile’ and become spread out. The question arises whether meerkat pups that experience different constraints to those facing chicks have evolved similar begging strategies. This paper describes the vocalisations that meerkat pups emit in the context of begging and investigates the influence of these calls on food allocation by older group members and on the behaviour of littermates. Meerkat pups use two types of calls when soliciting food from a potential feeder. The most common is a ’repeat’ call, which pups emit continuously when following an older forager over several hours a day. In addition, when a potential feeder finds a prey item, the pups next to it emit a bout of calls with increased calling rate, amplitude and fundamental frequency, termed ’high-pitched’ calls. Observations, together with playback experiments, showed that more prey was allocated to pups that called longer and more intensely. The pup closest to a feeder was almost always fed. The probability of emitting high-pitched calls did not depend on the time since a pup had received food, and the change from repeat to high-pitched calls occurred suddenly. The main function of the high-pitched call, therefore, does not appear to be to signal a pup’s hunger state. More likely, the two calls, in the context of begging, may be an adaptation to energetic constraints in a mobile feeding system. Pups, which are dispersed during foraging, may emit repeat calls over long periods to prevent potential feeders from eating all the prey themselves. At the moment a potential feeder finds prey, pups may give the more intense high-pitched calls to direct feeders to bring the food item to them and not to a littermate. Therefore, unlike the stationary feeding system where chicks emit one type of begging call when the feeder approaches the nest, meerkats, with a mobile feeding system, have evolved two discrete types of vocalisations in the context of begging. Received: 22 November 1999 / Revised: 1 July 2000 / Accepted: 17 July 2000     

Begging affects parental effort in the pied flycatcher, Ficedula hypoleuca

It has been suggested that nestlings use begging to increase their share of parental resources at the expense of current or future siblings. There is ample evidence that siblings compete over food with nestmates by begging, but only short-term effects of begging on parental provisioning rates have been shown. In this study, we use a new experimental design to demonstrate that pied flycatcher (Ficedula hypoleuca) nestlings that beg more are able to increase parental provisioning rates over the major part of the nestling period, thus potentially competing with future siblings. Parents were marked with microchips so that additional begging sounds could be played back when one of the parents visited the nest. By playing back begging sounds consistently at either male or female visits, a sex difference in provisioning rate that lasted for the major part of the nestling period was induced. If each parent independently adjusts its effort to the begging intensity of nestlings, begging may also be the proximate control mechanism for the sexual division of labour. Received: 24 January 1997 / Accepted after revision: 30 August 1997     

Parent-absent begging in the Brown-headed Cowbird (<Emphasis Type="Italic">Molothrus ater</Emphasis>): the role of short-term need and nestmate size

Although it is well-established that nestlings of many altricial species beg when parents are away from the nest, we have a poor understanding of parent-absent begging in brood parasites, including the proximate factors that may influence begging frequency and intensity. In this study, I examined how parent-absent begging was influenced by competitive asymmetries between host and Brown-headed Cowbird (Molothrus ater) nestlings under disparate levels of short-term need. Food-deprived cowbird nestlings begged more frequently and for a greater proportion of parent-absent period than when food-supplemented, with similar patterns observed in hosts of different sizes. In contrast, three metrics of cowbird begging intensity varied relative to host size but not due to differences in short-term need. Cowbirds consistently begged more frequently and intensively than host nestlings for a given level of short-term need, providing evidence that cowbird begging displays are more frequent and intense than non-parasitic nestlings during both feeding visits and parent-absent periods. In sum, the frequency of begging by cowbirds was only influenced by short-term need, whereas begging intensity during parent-absent events was only influenced by the host against which cowbirds competed. This study demonstrates that host size and short-term need had differing influences on the frequency and intensity of parent-absent begging in cowbirds, although both factors are likely important in limiting the evolution of parent-absent begging in cowbirds. Because it appears to provide no immediate benefits yet may decrease fitness, parent-absent begging should be included in future theoretical models investigating the evolution of begging displays in nestling birds.     

Reproductive effort of colonial and solitary breeding tree sparrows Passer montanus L.

A total of 250 nestboxes were arranged in five plots in a suburban area of Budapest, Hungary (19°04E, 47°41N). In each plot, 25 were placed at 50 m intervals to simulate solitary breeding and 25 3–5 m apart to simulate colonial breeding. Length of nest building period, feeding frequency, nestling mortality, nestlings' diet, productivity and parental condition were compared for colonial and solitary breeding tree sparrows Passer montanus. Parents with long nest-building periods, including the majority of first-year females, produced fewer young than parents which built over short periods. Parents fed nestlings morefrequently and nestlings had lower mortality in second than first broods; whether or not a third brood was reared was determined by the costs invested in first and second broods. Females that laid a third clutch had reared fewer young in first and second broods and were heavier than females that reared many young in two broods. Colonial birds had higher feeding frequencies, more similar diets and suffered lower nestling mortality than solitary parents for first broods, but they fed less frequently, diets were less similar, and nestling mortality was higher in second and third broods. It is suggested that colonial breeders benefited from the social stimulation of simultaneous feeding in first broods, but the advantage of synchronicity in feeding declined in second broods and the sparser breeding spacing of solitary parents was more advantageous for feeding in second and third broods. Birds that changed nest spacing between broods had fed nestlings less frequently and had higher nestling mortality before changing than birds which retained their spacing. Parents which changed from colonies to solitary nests fed more frequently with lower nestling mortality in the next brood than parents which retained colonial nests for their second (and third) brood. Solitary parents did not show such a relationship. The rearing of three broods caused higher weight loss in colonial than solitary parents.Correspondence to: L. Sasvári     

Resource allocation varies with parental sex and brood size in the asynchronously hatching green-rumped parrotlet (<Emphasis Type="Italic">Forpus passerinus</Emphasis>)

When eggs hatch asynchronously, offspring arising from last-hatched eggs often exhibit a competitive disadvantage compared with their older, larger nestmates. Strong sibling competition might result in a pattern of resource allocation favoring larger nestlings, but active food allocation towards smaller offspring may compensate for the negative effects of asynchronous hatching. We examined patterns of resource allocation by green-rumped parrotlet parents to small and large broods under control and food-supplemented conditions. There was no difference between parents and among brood sizes in visit rate or number of feeds delivered, although females spent marginally more time in the nest than males. Both male and female parents preferentially fed offspring that had a higher begging effort than the remainder of the brood. Mean begging levels did not differ between small and large broods, but smaller offspring begged more than their older nestmates in large broods. Male parents fed small offspring less often in both brood sizes. Female parents fed offspring evenly in small broods, while in large broods they fed smaller offspring more frequently, with the exception of the very last hatched individual. These data suggest male parrotlets exhibit a feeding preference for larger offspring—possibly arising from the outcome of sibling competition—but that females practice active food allocation, particularly in larger brood sizes. These differential patterns of resource allocation between the sexes are consistent with other studies of parrots and may reflect some level of female compensation for the limitations imposed on smaller offspring by hatching asynchrony.     

Food supply differentially affects sibling negotiation and competition in the barn owl (Tyto alba)

In contrast to most birds, nestling barn owls (Tyto alba) vocalise not only when parents are at the nest but also in their absence. Calls produced in their absence have been shown to facilitate sibling negotiation over the impending food resource. Since nestlings vocalise more vigorously in the presence of parents, they may be calling not to negotiate resources but to compete amongst each other over parental food distribution. A critical issue is to determine whether offspring need differentially affects sibling negotiation and sibling competition, that is vocalisation in the absence and presence of parents. To answer this question, I manipulated the food supply of 26 broods by adding or removing prey items. In the absence of parents, food-added broods vocalised at a significantly lower level than food-removed ones. In contrast, once a parent arrived at the nest, the vocalisation level was not significantly related to the manipulation of brood food supply. This suggests that in the absence of parents, it is more important for food-removed nestlings to vocalise intensely, and that in their presence, the relationship between begging and offspring need tends to vanish. In other words, brood food supply may affect sibling negotiation to a larger extent than sibling competition.     

Parental favouritism strategies in the asynchronously hatching European Roller (<Emphasis Type="BoldItalic">Coracias garrulus</Emphasis>)

In altricial birds, resource allocation during early developmental stages is the result of an interaction between parental feeding decisions and scramble competition between nestmates. Hatching asynchrony in birds leads to a pronounced age hierarchy among their offspring. Therefore, whenever parents exert control over resource allocation parents feeding asynchronous broods should simultaneously assess individual offspring internal condition and age. In this study, we first studied whether the highly ultraviolet (UV) reflective body skin of nestlings in the asynchronous European Roller (Coracias garrulus; roller hereafter) relates to nestling quality. In a second stage, we experimentally studied parental biases in food allocation towards senior and junior sibling rollers in relation to a manipulation of UV reflectance of the skin of their offspring. Heavier roller nestlings had less brilliant and less UV saturated skins than weaker nestlings. In our experiment, we found that parents with large broods preferentially fed nestlings presenting skin coloration revealing small body size (i.e. control nestlings) over nestlings presenting skin coloration revealing large body size (i.e. UV-blocked nestlings). Within the brood, we found that parental food allocation strategy depended on nestling age: parents preferentially fed senior nestlings signalling small body size, but did not show preference between control and UV-blocked junior nestlings. These results emphasise that parent rollers use UV cues of offspring quality while balancing the age of their offspring to adjust their feeding strategies, and suggest that parents may adopt finely tuned strategies of control over resource allocation in asynchronous broods.     

Food abundance and parental care in yellow warblers (Dendroica petechia)

Emlen and Oring (1977) suggested that monogamy in birds is maintained because of the need for strict biparental care. A corollary of their suggestion is that paternal care should decrease under conditions of high food abundance. An alternative is that paternal care would increase if males take advantage of the higher food abundance by trying to reduce the length of the nestling feeding period. We tested these two ideas using yellow warblers (Dendroica petechia) by providing some pairs with supplemental food, thereby reducing the importance of biparental care. However, the extra food did not decrease paternal effort, nor did it increase it (Fig. 2). Early in the nestling period experimental females brooded more but visited their nestlings less than did control females, but later, when brooding times decreased, experimental females fed their nestlings more than did control females (Fig. 3). There were no significant differences in nestling survival (Fig. 5), but nestlings in the control treatment were larger and heavier up to 6 days old (Fig. 6). The main effect of supplemental food was on maternal, not paternal behaviour. Models of biparental care assume interdependence between the parental effort of both parents. In this species, however, males and females provide for their brood independently from each other.     

Offspring reproductive value and nest defense in the magpie (Pica pica)

Summary Magpie (Pica pica) brood defense against a human at the nest was studied in a Mediterranean population with low renesting potential. Variations in two defense measures recorded during 106 trials at 41 different nests were positively correlated with brood age. Ineremental effects due to the number of successive visits to nests by us, brood size, and the time in the breeding season were not significant. Partial correlation analyses showed that visit rate was not an important determinant of nest defense, which thus favors an adaptive explanation of nest defense patterns. Two functional hypotheses to account for the increase in defense intensity with brood age were tested: whether (1) increased parental defense serves to compensate the higher predation risk of older nests or (2) increased parental defense reflects the increasing reproductive value of nestlings as they grow older. Daily mortality and incidende of predation (estimated from contribution of whole-brood losses to total mortality) was higher early in the nestling period, hence providing weak evidence for the assumption on which hypothesis (1) is based. The timing of parental defense intensity did not mirror variations in predation risk for the nest but variations in reproductive value of the brood, as can be estimated from daily mortality, thus supporting hypothesis (2). Magpie parents increased defense intensity in response to premature escaping by almost fully-developed nestlings. Since such a response lowers predation risk for the offspring and increases their probability of survival, this finding supports hypothesis (2), but runs contrary to hypothesis (1). Parents also increased defense in response to play-backs of alarm calls uttered by nestlings during escaping episodes. It is argued that parents should continuously monitor the degree of offspring development in order to assess their reproductive value and that, by alarm calling, chicks honestly make their parents aware of the gain in reproductive value that results from enhancement in locomotory abilities that occur at the end of the nestling period.     

Brood parasitism disproportionately increases nest provisioning and helper recruitment in a cooperatively breeding bird

Obligate avian brood parasites lay their eggs in nests of other species (hosts), which raise parasitic young. Parasitic nestlings are likely to influence host’s parental behaviours as they typically beg for food more vigorously than young host for a given hunger level. However, few studies have tested this idea, with conflicting results. These prior studies were largely limited to biparental hosts, but little is known about the effect of brood parasitism on parental behaviours in hosts that breed cooperatively. We followed a multimodel approach to examine the effect of brood parasitism on nest provisioning and helper recruitment in the baywing (Agelaioides badius), a cooperative breeder parasitised by screaming (Molothrus rufoaxillaris) and shiny (Molothrus bonariensis) cowbirds. Multimodel inference results indicated that feeding visits increased with nestling age, cooperative group size and number of cowbird nestlings in the brood. Brood size had little influence on feeding visits, which further suggests that baywings adjusted their provisioning effort in response to cowbird parasitism. In addition, nests parasitised artificially with shiny cowbird eggs or hatchlings recruited more helpers than unmanipulated nests having only host or screaming cowbird young. Our results provide novel evidence that brood parasitism and cooperative breeding interact in determining the levels of nest provisioning.     

Great spotted cuckoo fledglings are disadvantaged by magpie host parents when reared together with magpie nestlings

The post-fledging period is a critical phase for juvenile survival, and parental care provided during this period is a key component of avian reproductive performance. Very little is known about the relationships between foster parents and fledglings of brood parasites. Here, we present the results of a 5-year study about the relationships between fledglings of the non-evictor brood parasitic great spotted cuckoo (Clamator glandarius) and its magpie (Pica pica) foster parents. Sometimes, great spotted cuckoo and magpie nestlings from the same nest can fledge successfully, but most often parasitic nestlings outcompete host nestlings and only cuckoos leave the nest. We have studied several aspects of cuckoo post-fledging performance (i.e. feeding behaviour, parental defence and fledgling survival) in experimental nests in which only cuckoos or both magpie and cuckoo nestlings survived until leaving the nest. The results indicate that great spotted cuckoo fledglings reared in mixed broods together with magpie nestlings were disadvantaged by magpie adults with respect to feeding patterns. Fledgling cuckoos reared in mixed broods were fed less frequently than those reared in only cuckoo broods, and magpie adults approached less frequently to feed cuckoos from mixed broods than cuckoos from only cuckoo broods. These results imply that the presence of host's own nestlings for comparison may be a crucial clue favouring the evolution of fledgling discrimination; and furthermore, that the risk of discrimination at the fledgling stage probably is an important selection pressure driving the evolution of the arms race between brood parasites and their hosts.     

Feeding frequency and parental division of labour in the double-brooded great tit Parus major

Summary We studied the relative contribution of each sex and total effort expended in feeding nestlings in the great tit Parus major in relation to artificially altered brood size. A recent model suggests that feeding frequency should reflect the optimal trade-off between parental and fledgling survival, the former being negatively, the latter positively, influenced by high feeding frequencies. In both sexes weight loss was linearly related to feeding frequency. Since fledgling survival increases with nestling weight, the conditions of this model are fulfilled. However, in contrast to the predictions of the model, the total feeding frequency for both sexes combined did not differ between control and enlarged broods, but was lower for reduced ones. This outcome was not the result of a physiologically related inability of the parents to increase their delivery rate. Instead, we suggest that parents with enlarged broods could not find sufficient amounts of prey large enough to be economically worth transporting to the nest. Differences in brood-provisioning rates between the sexes may arise because costs and benefits of feeding nestlings may differ. Females lost more weight than males during the nesting period, but maintained a relatively higher weight during the incubation period. The relationship between weight loss and feeding frequency was similar for both sexes. Male and female brood-feeding frequency was related to brood size in a similar way. This is discussed in light of the great tit's mating system and the fact that the great tit is facultatively double-brooded.     

An experimental study of paternal behavior in red-winged blackbirds

Summary The effect of brood size and female nesting status on male parental behavior was investigated in red-winged blackbirds Agelaius phoeniceus using brood size manipulation experiments. Male redwings allocated parental effort on the basis of brood size and nestling age. Males began assisting females only at nests with at least three offspring older than three days. Female nesting status had no singificant influence on male parental care. When females were unable to meet a brood's demand for food, males assisted females with nestling feeding. Females did not reduce the amount of food delivered to nestlings when males assisted. The amount of food brought to nestlings by the male was additional to the amount of food provided by the female. Male assistance increased fledgling success. When female provisioning was sufficient to meet a brood's demand for food males did not assist. The value of male parental care varied inversely with the ability of the female to meet nestling food demands. The ability of unassisted females to provide sufficient food and to raise a brood of nestlings successfully appeared to be influenced by resource abundance.