Begging and short-term need in cowbird nestlings: how different are brood parasites?

Because brood parasitic nestlings are usually unrelated to their nestmates and to the provisioning adult, they are free from indirect costs of begging. Consequently, they are predicted to beg more intensely than host nestlings, and some models predict they will beg at an invariantly high level, regardless of short-term need. Previous work has shown intense begging by parasitic cowbirds, but short-term need was not controlled. In this study, we manipulated short-term need and measured begging intensity in two species pairs of host and parasitic nestlings: shiny cowbirds (Molothrus bonariensis) parasitizing larger hosts than themselves (rufous-bellied thrushes, Turdus rufiventris), and brown-headed cowbirds (M. ater) parasitizing smaller hosts than themselves (yellow warblers, Dendroica petechia). All four species increased their begging intensity with short-term need, though the change was much less pronounced between food-deprived and control treatments than between control and hand-feeding treatments. Shiny cowbirds begged more intensely than rufous-bellied thrushes following each treatment. In contrast, brown-headed cowbirds did not beg significantly more intensely than yellow warblers under any of the treatments. Intense begging by both species of parasites was as effective as host begging in stimulating the adults to make provisioning visits, but shiny cowbirds were less successful at acquiring food from adult thrushes. A wide array of factors may underlie the apparent differences in the begging behavior and success of brown-headed and shiny cowbirds, including relative size, experience, and local risk of nest predation. Our experiments clearly demonstrate, however, that these two species of parasitic cowbirds adjust their begging intensity based on short-term need.Communicated by H. Kokko     

Begging behavior and food acquisition by brown-headed cowbird nestlings

Understanding the selective forces that limit the exaggeration of begging signals is a critical issue in understanding the evolution of begging behavior. I studied the begging behavior of nestlings of the brown-headed cowbird (Molothrus ater), a brood parasite. In the nests of indigo buntings (Passerina cyanea), brown-headed cowbird nestlings received approximately twice as much food per hour than their host nestmates. I tested three hypotheses for the mechanism by which cowbirds acquired more food than their bunting nestmates: the size advantage hypothesis, the signal exaggeration hypothesis, and the novel begging behavior hypothesis. I found support for the hypotheses that cowbirds acquire more food as a result of their larger body size, and due to the exaggeration of begging signals that are not dependent on body size. I did not find support for the role of novel begging behaviors in cowbird food acquisition. These results suggest that food acquisition by host chicks in unparasitized nests could be increased by the exaggeration of begging signals. Recent work suggests that such exaggeration may be limited by the risk of nest predation, but further studies are needed. Received: 12 December 1997 / Accepted after revision: 29 December 1997     

Sex-related asymmetry in competitive ability of sexually monomorphic barn swallow nestlings

In diverse taxa, offspring solicit parental care using complex displays, which may evolve as reliable signals of condition or as mechanisms to manipulate parental investment. Differential sex allocation may therefore result from adaptive parental decisions or sex-related variation in competitive ability or because of sex-related asymmetries in kin selection. Under normal food provisioning, female barn swallow (Hirundo rustica) nestlings begged more loudly but did not receive more food than male nestlings. After food deprivation, begging call loudness of males but not females increased. Begging loudness positively predicted the number of feedings received by the nestlings, and males gained more mass than females after food deprivation. Male nestlings are more severely affected by chronic food reduction and may therefore accrue a larger benefit compared to females by increasing their food intake under short-term conditions of food scarcity. These results suggest that either females do not increase begging intensity to favour male broodmates which are more vulnerable to prolonged food stress, or that males prevail in scramble competition despite being similar in size to females.     

Patterns of egg yolk antioxidant co-variation in an avian brood parasite–host system

Despite their importance to offspring fitness outcomes, there has been little previous work on egg-mediated maternal effects in avian brood parasites which lay their eggs in the nests of other species. Here, we examine patterns of egg yolk antioxidant deposition in an avian host (red-winged blackbird, Agelaius phoeniceus) and their brood parasite (brown-headed cowbird, Molothrus ater). Cowbird nestlings compete directly for food with host blackbird chicks and succeed, despite hatching from a smaller egg, by begging more intensely and growing as or more rapidly than their hosts. Dietary-derived antioxidants, such as carotenoids and vitamins E and A, deposited in egg yolk, may enhance growth rate and reduce oxidative stress, and therefore provide a potential mechanism by which female cowbirds could enhance the competitiveness of their young in host nests. However, provision of antioxidants to eggs is costly so we predicted that female cowbirds should adjust the amount of antioxidants in response to variation in host quality. We found that whilst red-winged blackbird clutches parasitised by cowbirds did not differ in the pattern of antioxidant deposition compared to non-parasitised clutches, levels of vitamin E deposited in cowbird eggs were closely matched to that of the individual host clutch in which cowbirds laid their egg. This supports the prediction that female cowbirds adjust the antioxidant concentration of their eggs to increase the likely competitiveness of their offspring in the host nest into which they are laid and may help explain the success of cowbirds in competing with larger host young.     

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     

Begging and provisioning in broods of asynchronously-hatched yellow-headed blackbird nestlings

Studies of begging have found a positive relationship between begging level and provisioning level. Studies of unequal nestlings, however, have found that small nestlings generally beg more but are fed less than their larger siblings. We manipulated the begging levels of yellow-headed blackbird (Xanthocephalus xanthocephalus) chicks to investigate how begging benefits individuals in broods of unequal siblings. Food-deprived chicks begged more and were fed more; satiated chicks begged less and were fed less. When we deprived each chick of a brood in turn, large and small chicks generally increased begging and received more provisioning. Small chicks, however, rarely received more food than their larger siblings even when they behged relatively more. Parent yellow-headed blackbirds increase provisioning to hungry begging chicks, but also allocate food based on relative offspring size.     

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.     

Honest begging in the blue-footed booby: signaling food deprivation and body condition

To determine whether solicitation by blue-footed booby chicks accurately encodes their need for food, we independently manipulated the body condition and recent food ingestion of singleton chicks and recorded three measures of begging. Variations in the three measures of begging covaried only partially, but in general, chicks begged more intensely when they were in poor body condition and also when suffering recent food deprivation. Effects of body condition and recent food deprivation on begging were broadly additive, although deprived chicks in poor condition failed to beg more intensely than those in good condition. Protracted short-term deprivation may create such a high level of need that the body condition component of need becomes temporarily unimportant. Parents more frequently fed chicks that begged more intensely, chicks in poor condition, and chicks suffering food deprivation. Deprived chicks in poor condition received more food than deprived chicks in normal condition even though they did not beg more intensely, possibly because parents responded not only to current begging but also to begging earlier in the day, or to other cues to body condition. These results support the hypothesis that begging of boobies represents honest signaling of need.     

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.     

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 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.     

Maternal effects on begging behaviour: an experimental demonstration of the effects of laying sequence, hatch order, nestling sex and brood size

Differential resource allocation by females across the laying sequence has been hypothesised as a mechanism through which females could either compensate nestlings that hatch last in asynchronous broods or promote brood reduction. In this study we artificially incubated eggs and cross-fostered offspring to manipulate nestlings’ position in the hatching order, to identify whether the competitive ability of nestlings is dependent on position in the laying sequence. In both control and experimentally reversed broods, first hatched chicks had a higher survival than last hatched siblings. Yet, nestlings that hatched from eggs laid in the second half of a clutch begged with a greater intensity than nestlings hatched from eggs laid in the first half of a clutch. In natural broods, the greater begging competitiveness of nestlings from later-laid eggs led to a moderation of sibling competition and these nestlings achieved the same body size and weight as nestlings from eggs laid in the first half of the clutch. The lack of a substantial difference in the size and condition of surviving nestlings in respect to laying order suggests that differential resource allocation across the egg-laying sequence partially compensates for hatching last in asynchronous broods and reduces the negative effects of the nestling size hierarchy. The effect of laying order, brood size and experimental treatment also differed for male and female nestlings. Our study highlights the need to be aware of the complex and subtle effects of nestling sex and laying sequence when investigating genetic and environmental influences on individual fitness.     

Differential parasitism of least flycatchers and yellow warblers by the brown-headed cowbird

Summary Brown-headed cowbirds (Molothrus ater) parasitized yellow warblers (Dendroica petechia) six times more frequently than least flycatchers (Empidonax minimus) nesting in the same riparian habitat in southern Manitoba. This difference was surprising because least flycatchers were higher quality hosts; flycatchers always accepted cowbird eggs while warblers did so on only about half the occasions. Both hosts were equally good foster parents, at least until fledging; thus, once an egg was accepted there was probably no further advantage in parasitizing one species over the other. The probability that a nest was parasitized decreased with increasing nest height, perhaps as a consequence of the cowbird's habit of searching for nests close to the ground. Since least flycatchers nested higher in the canopy than yellow warblers, a greater proportion of flycatcher nests probably were not detected by cowbirds. Nevertheless, nest height alone could not account fully for the lower incidence of parasitism on flycatchers. Clutch initiation in flycatchers peaked 6 days after warbler clutch initiations and 5 days after that of cowbirds. Despite the later peak in initiations, flycatcher nests were always available as potential hosts over the entire cowbird laying season and it was not until new clutch initiations by warblers declined in late summer that flycatchers were actually used as hosts. Because least flycatchers responded more aggressively than yellow warblers to a model female cowbird at the nest, we concluded that greater nest defense by flycatchers may have also reduced the rate of brood parasitism in this host. Together, our results suggest the large difference in the frequency of parasitism between these two hosts was primarily a product of nest location but that differences in host nest-defense and breeding season asynchrony may have contributed to preferential host selection.Offprint requests to: J.V. Briskie     

An experimental study of competition for food between male and female nestlings of the red-winged blackbird

Summary Ten experimental broods of red-winged black-bird (Agelaius phoeniceus) nestlings, experimentally manipulated to contain two males and two females of similar age, were observed with the aid of video cameras to determine (a) which nestling characteristics were most important in influencing its chances of being fed and (b) if males and females differed with respect to these characteristics. Nestlings that could reach highest while begging were most successful at obtaining food from parents during individual feeding bouts. However, while there was a tendency for nestlings that begged earlier to be offered food first, a nestling's position in the nest during these bouts did not influence its feeding success. Males begged more than often females and were offered more food by parents. Males were also more likely to be fed when they begged. This was most likely because males, being larger than females, were able to reach higher while begging. Neither sex was more likely to beg sooner or occupy certain positions in the nest. However, while males that had been transferred from other nests received less food than natal males, there were no differences in food acquisition between transferred and natal females. Differences in the relative success of males and females under different conditions may help explain differences in fledgling sex ratios observed in many dimorphic species.     

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     

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.     

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     

Solicitation displays reliably reflect oxidative damage in barn swallow nestlings

Reproduction entails parental decisions over allocation of limiting resources. ‘Begging’ solicitation displays by the offspring have been hypothesised to evolve as reliable indicators of quality, allowing parents to adopt optimal allocation strategies. To function as such, however, begging signals must covary with offspring traits that affect viability. Here we tested whether oxidative damage of nestling barn swallows predicts begging bout duration, call amplitude and intensity of postural displays both under normal food provisioning and under food shortage. Oxidative damage as determined by normal metabolism, pathological conditions and functioning of the immune system can serve as a comprehensive indicator of offspring condition because it can profoundly affect viability, senescence and reproduction. Begging bout duration was negatively predicted by oxidative damage but only after food deprivation. Postural begging negatively covaried with oxidative damage, at least when nestlings were confronted with related nest mates. Plasma antioxidant capacity did not predict begging behaviour. Thus, we show for the first time that behavioural begging covaries with a comprehensive index of chick condition, lending support to reliability of such begging traits also as signals of quality rather than need or hunger only. Future studies are candidate to decouple the effect of hunger state and individual condition on behavioural begging.     

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.     

Spectral tuning and perceptual differences do not explain the rejection of brood parasitic eggs by American robins (Turdus migratorius)

By laying their eggs in the nests of other birds, avian brood parasites impose the cost of rearing young upon their hosts. The recognition and rejection of foreign eggs are primary host defenses against costly brood parasitism. Hosts of parasitic brown-headed cowbirds (Molothrus ater) challenge coevolutionary theory because most cowbird hosts accept parasitic eggs despite their drastically different appearance from the hosts’ own eggs. American robins (Turdus migratorius) are one of only 10 % of the over 200 potential cowbird host species to robustly reject parasitic eggs, but the mechanisms driving the sensory bases of foreign egg rejection remain elusive. Our research combined avian visual perceptual modeling and behavioral experimentation to investigate chromatic cues eliciting parasitic egg rejection in American robins. We assessed the effects of perceivable background color differences between real host and model parasite eggs, across all four avian photoreceptors, on rates of rejection of model eggs spanning in color across the entire avian spectral sensitivity range, and including immaculate model eggs matching the natural colors of robin and cowbird eggs. The results suggest that egg rejection in robins is driven by the overall perceivable difference in color between own and artificial eggs, and input from all four single-cone avian photoreceptors affects the rejection decision. The results, however, also reveal that when viewed by the avian eye, natural cowbird eggs appear more similar in background color to robin eggs than predicted by the high rejection rate of these parasitic eggs. This suggests that robins respond specifically to parasitism by cowbirds, despite an apparent lack of sensory tuning toward the detection of the background color of cowbird eggs.