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.     

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     

Sibling competition and cooperation in mammals: challenges,developments and prospects

Many vertebrates grow up in the company of same or different-age siblings, and relations among them can be expected to significantly influence individual life histories and the development of individual morphological, physiological, and behavioral phenotypes. Although studies in birds still dominate and have stimulated most theoretical considerations, the increasing number of mammalian studies promises to broaden our understanding of this complex field by enabling interesting comparisons with the rather different bird system. It therefore seems timely to bring together recent studies of sibling relations in mammals and to demonstrate what these can offer in the way of fresh insights. In this brief review, intended to accompany a series of papers on a diverse range of mammals, we outline the current state of sibling research in mammals, comparing it to the better studied birds. Most obviously, in mammals, mother and young are in much closer contact during early life than in birds, and siblings can influence each other’s development as well as the mother’s physiology while still in utero. During nursing, mammalian young also encounter a very different feeding situation to bird siblings. These contrasts should help stimulate further debate, as well as provide further opportunities to study the relative importance of maternal versus sibling effects on individual development. Finally, we discuss the need to balance studies of sibling competition and conflict with a consideration of the benefits accruing to individuals from sibling presence and the need for long-term studies of the influence of early sibling relations on individual development and life histories. This contribution represents the introduction to the special issue “Sibling competition and cooperation in mammals”.     

No reduction in aggression after loss of a broodmate: a test of the brood size hypothesis

In some vertebrate species, parents create a large brood or litter then, in the event of unfavourable ecological conditions, apparently allow the number of offspring to be adaptively reduced through siblicide. But how is sibling aggression regulated so that deaths occur only in unfavourable conditions? One proposed mechanism is brood size-dependent aggression. Two experiments tested for this mechanism by reducing three-chick broods of blue-footed boobies either during or after the period of dominance hierarchy establishment. In neither experiment did aggression of the two eldest and highest ranking chicks decline after removal of the youngest broodmate, in comparison with controls. These results suggest that dominant booby chicks do not become less aggressive to each other after disappearance of their youngest broodmate and that this species does not show brood size dependent aggression. Elder blue-footed booby chicks increase their attacks on broodmates when they receive less food, and this mechanism may be sufficient to tailor brood size to food availability.     

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.     

Learning during competitive positioning in the nest: do nestlings use ideal free ‘foraging’ tactics?

Begging behaviour of nestling birds may involve more than a simple, honest source of information for parents to use in provisioning. Many aspects of begging behaviour relate instead to sibling competition for food items within the nest, and we might expect evidence of adaptive learning and behavioural adjustment in response to experience of the competitive environment. In this study, we consider begging in different locations within the nest as analogous to foraging in different patches, varying in food availability. Using hand-feeding trials, we created zones of differing profitability within an artificial nest by adjusting either the prey size or number of items delivered, and allowed only indirect competition between pairs of southern grey shrike (Lanius meridionalis) nestlings. Nestlings demonstrated the ability to detect differences in zone profitability and position themselves accordingly. By the end of both the prey size and delivery rate trials nestlings had increased the amount of time spent in the high quality zone. Such movement in response to differences in load quality, as well as frequency, demonstrates the ability of nestlings to learn about their environment and to facultatively adjust their begging in order to maximise energetic rewards.     

Ladybird mothers mitigate offspring starvation risk by laying trophic eggs

A large proportion of ladybird beetle (Coleoptera: Coccinellidae) eggs are apparently infertile—they do not develop an embryo and are consumed by larvae hatching within the egg batch. The predicted benefits of egg consumption for larvae are empirically well supported. An important question, however, remains: are these eggs a maternal strategy to feed offspring (i.e., trophic eggs) or did egg eating evolve to exploit unavoidably infertile eggs? We investigated the adaptive value of infertile eggs in laboratory experiments with multicoloured Asian ladybirds (Harmonia axyridis). Female H. axyridis were assigned to low and high resource environments for brief intervals; we predicted that tactics to facilitate egg cannibalism, such as infertile egg production and hatching asynchrony, would be adopted in low food environments in which starvation risk for offspring is greater. We conducted two experiments in this manner that provided females with information about resource levels through prey feeding or scent. We also observed female oviposition patterns and tested for infertile egg distributions that departed from random. Females produced 56% more infertile eggs in the low vs. the high food treatment; however, hatching synchrony did not change. We consider a potential confound between information and nutrition state unlikely because ladybirds are well able to tolerate low food for 24 h, the duration of trials, and because females were in good condition when trials began. Results suggest that ladybirds use information from prey encounter to manipulate the proportion of trophic eggs in a manner consistent with the adaptive hypothesis, the first evidence of trophic egg plasticity in a non-eusocial insect.     

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     

Effects of territory quality, food availability and sibling competition on the fledging success of oystercatchers (Haematopus ostralegus)

We investigated the fledging probability of oystercatcher, Haematopus ostralegus, chicks as a function of hatching order, brood size, territory quality and food availability. Sibling dominance was related to the hatching order in both low- (’leapfrogs’) and high-quality (’residents’) territories. Differences in hatchling mass might have aided the establishment of a dominance hierarchy, since breeders produced small late eggs and hatchlings. These mass differences were most pronounced in leapfrogs, and in large broods in years with lower food availability (’poor’ years). Late hatchlings fledged less often and with lower body masses compared to early hatchlings in all situations. Leapfrogs produced smaller broods and hatched their broods more asynchronously in poor years than leapfrogs breeding in years with more available food (’good’ years) and residents breeding in both poor and good years. Large brood sizes resulted in lower survival of hatchlings in poor years. These results favour the ’brood reduction’ hypothesis. However, contrary to the expectations of this hypothesis, hatching order also affected fledging success in residents. Moreover, large brood size resulted in higher survival of hatchlings in good years, particularly in residents. Thus, although large broods experienced losses due to sibling competition in some years, they nevertheless consistently produced more fledglings per brood in all years, both as leapfrogs and residents. We believe this effect is due to parental quality correlating with initial brood size. Most leapfrogs, at best, fledged one chick successfully each year, losing chicks due to starvation. Nevertheless, leapfrog broods were reduced in size after hatching significantly less quickly than resident broods. These results suggest that breeders lay and hatch insurance eggs to compensate for unpredictable losses due to the high predation rates on both nests (ca 50%) and chicks (ca 90%), in accordance with the ’nest failure’ hypothesis. Received: 14 February 2000 / Revised: 27 September 2000 / Accepted: 10 June 2000     

Parent–offspring resource allocation in domestic pigs

Behavioural research on domestic pigs has included parent–offspring conflict, sibling competition, and the use of signals which influence resource allocation. In this paper, we review key sow–piglet behavioural studies and discuss their relevance to resource allocation theory. Sibling competition begins in the uterus and continues after birth, as piglets compete directly for access to the sow’s teats. This competition is made more severe by a unique dentition, which newborn piglets use to lacerate the faces of siblings during teat disputes. Competition often leads to the death of some littermates, especially those of low birth weight. Piglets also compete indirectly for milk, apparently by stimulating milk production at the teats that they habitually use at the expense of milk production by other teats. The complex nursing behavior of the sow appears designed to prevent the more vigorous piglets from monopolizing the milk. Sows give vocal signals which both attract piglets to suckle and synchronize their behavior during nursing episodes. Piglets give loud vocal signals when separated from the sow; calls which vary in intensity and appear to be honest signals of need. Udder massage by piglets may also serve as an honest signal of need. Parent–offspring conflict has been demonstrated experimentally in pigs. Specifically, when given the opportunity to control contact with their piglets, sows nurse less frequently, provide less milk, and lose less weight during lactation than sows that cannot control their level of contact. Because of this interesting natural history and because they are so amenable to experimentation, domestic pigs provide a rich system for testing ideas drawn from resource allocation theory. This contribution is part of the special issue “Sibling competition and cooperation in mammals” (guest editors: Robyn Hudson and Fritz Trillmich).