Preferential allocation of food by magpies Pica pica to great spotted cuckoo Clamator glandarius chicks

Adult magpies Pica pica provide parasitic great spotted cuckoo Clamator glandarius nestlings with a diet very similar to that fed to their own chicks. In both naturally and experimentally parasitized nests, great spotted cuckoo chicks were fed at a higher rate than magpie chicks in the same nest. This preferential allocation of food by magpie parents to great spotted cuckoo chicks is consistent with the supernormal stimulus hypothesis, because this result implies that cuckoo chicks provide stronger stimuli for parental care than host chicks. Great spotted cuckoo chicks receive most of the food brought to the nest by the foster parents, because they exploit a series of stimuli which jointly (or sometimes individually) operate as a supernormal stimulus. This hypothesis predicts that if any stimulus is masked, the efficiency of the cuckoo in eliciting parental care will decrease. Here, we analyze experimentally the effects of two of these stimuli, preferential feeding of large nestlings and of nestlings with conspicuous palatal papillae. Firstly, when we experimentally introduced one medium-sized (7–9 days) cuckoo chick into an unparasitized magpie nest where the largest magpie chick was 12–15 days old, the cuckoo did not receive significantly more food than the average or the largest magpie chick. Secondly, when unparasitized nests were experimentally parasitized with a cuckoo chick that had its gape painted to mimic that of magpie chicks, the parasitic cuckoo received less food than the average magpie chick.     

Does the great spotted cuckoo choose magpie hosts according to their parenting ability?

When brood parasites are about to lay an egg, they have to decide which nest to parasitize. The best nest in which to lay will depend on the parenting ability of the host. We have studied selection of magpie (Pica pica) hosts by great spotted cuckoos (Clamator glandarius). Great spotted cuckoos preferentially parasitize large host nests. Nest volume in magpies is a good indicator of territory quality, since there is a negative relationship between magpie nest size and breeding date, and timing of breeding in magpies is known to be positively related to territory quality. Moreover, magpies occupying high-quality territories have high breeding success. Therefore, nest size is positively related to the quality of magpies. Parasitized magpie nests were of greater volume than the nearest neighbouring nest not parasitized by the great spotted cuckoo. In order to test whether the great spotted cuckoos might select high-quality magpie hosts, we manipulated pairs of parasitized and non-parasitized nests with identical laying dates and habitats, introducing into each of the nests the same number of parasitic and non-parasitic eggs. The number of fledglings reared (magpie plus great spotted cuckoo chicks) in naturally parasitized nests was higher than in experimentally parasitized nests. Thus, the probability of survival of the parasite chicks increased if cuckoo eggs were laid in the nests of high-quality hosts originally chosen by the parasite.     

Micro-evolutionary change in host response to a brood parasite

The relationship between brood parasites and their hosts is usually assumed to result in coevolution, and documentation of changes in extant populations should thus be possible. Here we describe how the ejection rate of eggs of an obligate brood parasite, the great spotted cuckoo Clamator glandarius, by its host, the magpie Pica pica, has recently increased in an area in southern Spain. The ejection rate of great spotted cuckoo eggs in naturally parasitized nests of the magpie increased at a rate of 0.5% year' during the period 1982–1992. This result was verified in a number of field experiments using nonmimetic and mimetic model eggs. The rate of increase in ejection rate was 4.7% year^-1 for mimetic eggs and 2.3% year^-1 for nonmimetic eggs. There were clear differences in parasitism by the great spotted cuckoo between study plots and years, which makes comparisons of rates of parasitism between areas difficult without considering temporal variation. The recent increase in the ejection response of magpies to great spotted cuckoo eggs was not due to magpies using the abundance of cuckoos as a cue to the intensity of parasitism.     

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.     

Brood mate eviction or brood mate acceptance by brood parasitic nestlings? An experimental study with the non-evictor great spotted cuckoo and its magpie host

Some avian brood parasitic nestlings are highly virulent, destroying all host eggs or nestmates, while others accept growing up together with host nestmates. The traditional idea was that all brood parasitic nestlings would benefit from being alone in the host nest. Thus, why do nestlings of some brood parasitic species accept the company of host offspring in the nest? The trade-off hypothesis suggests that brood parasites must balance the costs and benefits of killing host young because of two major potential costs: risk of nest desertion and loss of begging assistance. Here, we test this hypothesis in a non-evictor cuckoo species, the great spotted cuckoo Clamator glandarius and its main host, the magpie Pica pica, by manipulating brood size (1–3 nestlings) and brood composition (only cuckoo, only magpie or mixed) during three consecutive breeding seasons. None of the broods were abandoned by host parents, and cuckoo nestlings alone in the nest tended to grow faster (i.e. wing length). Thus, none of the predictions of the two potential costs on which the trade-off hypothesis is based apply to the great spotted cuckoo–magpie system. Our experimental study could not directly test why chick killing has not evolved in great spotted cuckoos, but the results point in the direction of several possibilities. We suggest that chick killing in great spotted cuckoos may not be adaptive mainly because another, less costly strategy (i.e. outcompeting host nestmates for food), is efficient for successful parasitism of magpie hosts.     

Location of suitable nests by great spotted cuckoos: an empirical and experimental study

Brood parasites depend entirely on their host species to raise their nestlings until independence. Thus, brood-parasite females must discover and select nests that are at a suitable stage for parasitism, and thus, the location of each parasitic egg is crucial in determining the brood-parasite female’s fitness. In relation to host behaviour, one of the main hypotheses proposed to explain how brood-parasite females find and select a suitable nest posits that the most active hosts during nest building should undergo a higher risk of being parasitised (the “host-activity hypothesis”). Here, using the great spotted cuckoo, Clamator glandarius–magpie, Pica pica system, we found that not only cuckoo females parasitise magpie nests regardless of the location and characteristics of nests, but also that the parasite’s observation of host activity near the nest determines a cuckoo female’s decision of laying in a nest. Only one experimental nest (without host activity) was parasitised before the magpie started laying, while 34.14?% of natural active nests were parasitised before the magpie started laying. These observations support the host-activity hypothesis for nest location in great spotted cuckoos.     

A nesting association between semi-colonial Bullock's orioles and yellow-billed magpies: evidence for the predator protection hypothesis

Mixed-species nesting associations have been reported for many birds. However, few studies have shown that one species specifically chooses to nest close to another in order to obtain a direct benefit and only two studies have managed to adequately test the predator protection hypothesis for such associations. Here we demonstrate a nesting association between Bullock's orioles (Icterus galbula bullockii) and yellow-billed magpies (Pica nuttalli). The evidence indicates that this is an active association in which, given a choice of suitable habitat areas in which to nest, orioles choose those areas that contain magpie nests. Oriole nests associated with magpie nests have a significantly reduced predation rate compared to those without associated magpie nests. This suggests that orioles prefer to nest near to magpies and that this preference has evolved due to the protection provided by neighbouring magpies. The nesting association results in the clumping of oriole nests around magpie nests, and therefore provides a possible explanation for the semi-colonial nature of Bullock's orioles. Received: 23 November 1998 / Received in revised form: 25 May 1999 / Accepted: 30 May 1999     

The multidimensionality of behavioural defences against brood parasites: evidence for a behavioural syndrome in magpies?

Studies of antiparasite defences against cuckoo parasites have largely neglected the possibility that behavioural components of host defence may correlate giving rise to a behavioural syndrome. Furthermore, the different contribution of the host’s sex in nest defence has traditionally been disregarded. Here, we studied magpie (Pica pica) mobbing behaviour towards dummies of great spotted cuckoo (Clamator glandarius) and non-harmful hoopoes (Upupa epops) and egg rejection of parasite eggs in a population of colour-banded magpies. We predicted a positive correlation between the intensity of nest defence and egg rejection within each sex and that females respond more intensely than males to the threat of brood parasitism as they undertake incubation. Magpie males, but not females, defended their nests more intensely in those nests in which cuckoo model eggs were rejected. Individual magpies did significantly differ in their baseline level of nest attentiveness; however, there were no individual differences once pair identity was considered. Males and females defended their nests more intensely when it was exposed to the presence of a great spotted cuckoo dummy. Males, but not females, were more prone to appear at their nests, and females, but not males, were more prone to defend more intensely when their nests were challenged by a parasite threat. Our results thus agree with the view that mobbing behaviour and egg rejection in magpies may actually constitute a pseudosyndrome and highlight the necessity to integrate interindividual variation and the sex of the host in studies of the evolution of host defences.     

Cuckoo growth performance in parasitized and unused hosts: not only host size matters

The quality and quantity of food delivered to young are among the major determinants of fitness. A parental provisioning capacity is known to increase with body size. Therefore, brood parasitism provides an opportunity to test the effects of varying provisioning abilities of different-sized hosts on parasitic chick growth and fledging success. Knowledge of growth patterns of common cuckoo, Cuculus canorus, chicks in nests of common hosts is very poor. Moreover, no study to date has focused on any currently unused hosts (i.e., suitable cuckoo host species in which parasitism is currently rare or absent). Here, I compare the growth performance of cuckoo chicks in nests of a common host (the reed warbler, Acrocephalus scirpaceus) and two unparasitized hosts (the song thrush, Turdus philomelos, and the blackbird, Turdus merula). Parasitic chicks were sole occupants of the observed nests, thus eliminating the confounding effect of competition with host chicks. Experiments revealed striking differences in parasitic chick growth in the two closely related Turdus hosts. Cuckoo chicks cross-fostered to song thrush nests grew much quicker and attained much higher mass at fledging than those in nests of their common reed warbler host. Alternatively, parasitic chicks in blackbird nests grew poorly and did not survive until fledging. I discuss these observations with respect to host selection by parasitic cuckoos.     

Experimental support for the use of egg uniformity in parasite egg discrimination by cuckoo hosts

Common cuckoo (Cuculus canorus) parasitism drastically reduces the reproductive success of their hosts and selects for host discrimination of cuckoo eggs. In a second stage of anti-parasite adaptation, once cuckoos can lay eggs that mimic those of their hosts, a high uniformity of host egg appearance within a clutch may favour cuckoo egg discrimination. Comparative evidence provides indirect support for this hypothesis although experimental support is currently lacking. Here, we studied the effect of experimentally decreased uniformity of host egg appearance on cuckoo egg discrimination by great reed warbler (Acrocephalus arundinaceus) hosts in a population in which long-term cuckoo parasitism has led to high levels of cuckoo–host egg mimesis. We manipulated host clutch uniformity by adding extra spots to fresh host eggs just after they were laid. Rejection of non-mimetic experimental eggs added to these nests was compared with those in control nests in which uniformity was not altered. Previously, by over-painting real spots in a control group of nests, we showed a negligible effect of our paints on hosts’ perception of their eggs. We show that for the great reed warbler, non-mimetic experimental eggs were relatively more tolerated in experimental nests, i.e. with lower uniformity (40%) than in control nests (5%). This is the first experimental study, to our knowledge, which demonstrates a reduced discrimination of foreign eggs as a consequence of an increase of egg phenotypes variation perception in a cuckoo host.     

Does supernormal stimulus influence parental behaviour of the cuckoo's host?

The supernormal stimulus hypothesis (SSH) states that a cuckoo chick should obtain more parental care than host young by means of exaggerated sensory signals. We tested the SSH by comparing parental care by reed warblers at parasitized and non-parasitized nests. A comparison of feeding rates to parasite and host chicks of the same size showed that parasitized nests received more food than non-parasitized ones with one host chick. There was an interesting relationship between average prey length and the mass of a cuckoo chick: prey length first increased with chick mass, but decreased after the cuckoo chick outgrew the average-sized host brood (three to four young at fledging). This might be expected if fosterers reduced the selectivity of their foraging behaviour when trying to satisfy the supernormal food demands of the parasitic chick. This suggestion is supported by the finding that the relationship between nestling mass and proportion of less economical small prey is inverse to the relationship between nestling mass and prey size. These results suggest that the parental behaviour of reed warblers is adjusted by selection to the needs of an average-sized brood. The overall proportion of insect orders was significantly different between the parasitic and host chicks. This result probably reflects the opportunistic foraging habits of the host. The qualitative difference (proportion of insect orders) between host and cuckoo nestling diets is partly a by-product of unequal length distribution of members of different taxonomic groups. The results of this study are consistent with the SSH.     

Parental care of nestlings by male red-winged blackbirds

Summary Nestling feeding by males is less common among birds with polygynous mating systems than in monogamous species, because of the pronounced trade-off between parental behavior and the attraction of additional mates. In this study, however, we found that male red-winged blackbirds (Agelaius phoeniceus) commonly assisted females in feeding nestlings in several Ontario marshes. Male parental care was additional to that provided by females, and it significantly enhanced the fledging success of nests (Table 2). Male redwings did not help to feed all nests on their territories: primary and secondary nests were much more likely to receive male parental care than tertiary and later nests. Contrary to expectation, male parental care was not restricted to the nests of primary females: a greater proportion of secondary than primary nests were assisted (Tables 4a and b). The presence of new females settling on the territory at the same time that a resident female had nestlings, resulted in males deferring parental care until a later brood. This suggests that males trade off the recruitment of females against parental care to an existing brood. Although the number of nestlings fledging from a male's territory was strongly influenced by the number of females in the harem, males could additionally increase their reproductive success by feeding nestlings in one or more nests on their territories (Fig. 2). The reproductive success of females was significantly enhanced by male assistance in feeding nestlings (Table 3). However, those females not receiving male assistance on territories of feeding males did not suffer a significantly reduced reproductive success in comparison to females on territories of non-feeding males (Table 2). Males varied considerably in the quality of brood care given. We therefore suggest that the quality of male parental care may be a factor considered by females in choosing a breeding situation.     

Brood reduction in the American white pelican (Pelecanus erythrohynchos)

Summary American white pelicans (Pelecanus erythrorhynchos) breeding in colonies at East Shoal Lake, Manitoba, Canada exhibited a mean hatching asynchrony of 2.5 days in 2-egg clutches. This resulted in a size difference between chicks which facilitated sibling dominance, harassment and lack of food for the subordinate chick. Only one young survived per nest. In marked broods, the secondhatched chick survived in 20% of successful nests. Manipulated clutch sizes (1, 2 and 3 eggs or chicks per nest) revealed that the presence of a second chick contributes significantly to the reproductive success of the parents. Results support the hypothesis that the second egg functions as a form of insurance against early loss of the first egg or chick. The parents, by establishing hatching asynchrony, by nonintervention in sibling aggression, and by selectively feeding the dominant chick, maximize their chance of rearing the most viable young.     

Help or disperse? Cooperation in termites influenced by food conditions

Ecological factors have been claimed paramount for the evolution and maintenance of cooperative group living in eusocial termites, as well as in cooperatively breeding birds and mammals. However, a clear demonstration of the role of any specific ecological factor in termites has been lacking. In the termite Cryptotermes secundus, individuals have two options, staying as helpers at the natal nest or developing into winged sexuals that disperse to found new colonies. An important ecological factor expected to influence the course of termite development is food availability; C. secundus nests inside a single piece of wood that serves as the sole source of food for the duration of the colony. As wood is consumed, the longevity of the colony is reduced, thus diminishing the potential fitness gains of staying at the nest. We experimentally investigated the occurrence of cooperative behavior and development under abundant- and limited-food conditions. Workers exposed to food-limited conditions were more likely to develop into dispersing sexuals and increased selfishly their food-acquisition behaviors. Proximately, a reduced frequency of proctodeal trophallaxis may have interfered with the distribution of pheromones that inhibit sexual development. Ultimately, decreased inclusive fitness benefits in food-limited, and thus short-lived nests, appear to explain the development of dispersing sexuals, supporting (1) the benefits-of-philopatry hypothesis as developed for the occurrence of cooperative breeding in vertebrates, and (2) predictions of reproductive skew theories.Communicated by L. Keller     

Mate choice in the yellow-shouldered widowbird: correlates of male attractiveness

Recent investigations of male ornaments in sexual selection have used experimental manipulation of tail length in three widowbird species, but only for one of these have correlates of male reproductive success been reported. I examined correlates of male attractiveness to nesting females over two breeding seasons for the polygynous yellow-shouldered widowbird, Euplectes macrourus, in order to discover which cues females may be using to select mates. The black, long-tailed ( 10 cm) males defend large territories and build nest frames, or cock's nests, which females then line and use for nesting. I examined various aspects of male morphology, five behavioral displays, territory characteristics, and the number of cock's nests that males built. Few correlates of mating success were found. The best predictor was the number of cock's nests that a male builds, though one courtship display also correlated with male mating success in 1 year, as did average grass height. Tail length did not correlate with male mating success. A partial correlation analysis confirmed that cock's nests and, in 1 year, grass height, were the primary contributers to male success. Females may choose where to nest primarily on the availability of suitable nesting sites. Long tails may be used by females seeking extra-pair copulations or in male-male competition for territories.     

Red-winged blackbird parental investment following brood parasitism by brown-headed cowbirds: is parentage important?

Parental investment by red-winged blackbirds (Agelaius phoeniceus) in response to natural and experi‐mental parasitism by brown-headed cowbirds (Molothrus ater), and in response to freeze-dried, female cowbird mounts presented near redwing nests during the egg-laying period was measured. Two measures of redwing parental investment were used: nest defense effort toward a model predator, and rate of feeding nestlings. There were no significant differences in levels of parental investment among unparasitized nests, naturally parasitized nests, or experimentally parasitized nests. Similarly, parental investment did not differ between redwings that were exposed to the cowbird mount and those that were not exposed to the mount, or among redwings exposed to the cowbird mount at different distances from the nest. This suggests that red-winged blackbirds do not recognize when they have been parasitized, and hence do not associate parasitism with a decrease in their parentage, or that parentage is not an important predictor of parental investment in this species. Received: 24 January 1997 / Accepted after revision: 7 June 1997     

Non-adaptive adoptions of nestlings in the colonial lesser kestrel: proximate causes and fitness consequences

We examined the causes, costs and benefits of adoption in the altricial lesser kestrel Falco naumanni. Specifically, we tested the intergenerational conflict hypothesis, proposed to explain adoption in some birds. Adoptions involved 76% of the nests and 51% of the nestlings at a mean age of 25 days (12 days before fledging). Nest-switching nestlings were not in poorer body condition, more parasitized or younger than their siblings, and body condition and prey delivery rates of their parents did not differ from those of other parents. In the foster nest, adopted nestlings did not benefit from higher feeding rates or a prolongation of the nestling period. They did not have fewer nest-mates or achieve higher rank within the new brood. Thus, adopted nestlings did not improve their body condition and survival. Adult lesser kestrels seemed unable to finely discriminate beween their own and alien chicks. Foster parents bore the cost of an increase of prey delivery rates, although it did not affect their survival or subsequent reproductive performance. Therefore, our results do not support the intergenerational conflict hypothesis, and suggest that adoption in this species is non-adaptive. Traditionally, the lesser kestrel bred in cliffs where movement among nest-sites was restricted. Nowadays, about half of the colonies are in tiled roofs which facilitate nest-switching by nestlings. The high rate of adoptions may thus be explained as reproductive errors associated with the recent occupation of a new breeding habitat. Received: 3 May 1996 / Accepted after revision: 19 January 1997     

Provisioning rules and chick competition in asynchronously hatching common terns (Sterna hirundo)

Interactions between nestling birds and their parents are models for examining parent–offspring communication and sibling competition. Most studies have focused on species where young are restricted to a nest. However, offspring of many species are mobile and fed by parents for an extended period post-hatch. These chicks mobility may provide an opportunity to examine the role of signalling and physical competition on parental feeding decisions. We examined parental provisioning rules in relation to offspring behaviour and hatching order (i.e., competitive ability) in a species with mobile young, the common tern. We determined that about 95% of feedings were directed to the first chick to reach the parent when it landed with food. We developed a probabilistic model to predict the likelihood of a chick reaching the parent first, and thus receiving food. Our model showed that begging intensity, feeding history, and the interaction between begging intensity and relative proximity to the parent best predicted which chick would arrive first. Increased begging was associated with arriving first significantly more when a chick was relatively further from the parent than when it was closer than its siblings. Independently of these factors, larger, earlier-hatched chicks were more likely to be fed than smaller, later-hatched chicks. Additional analyses showed that parents landed closer to more intensively begging chicks, however, increased begging did not explain the advantage of earlier-hatched chicks because begging intensity did not vary with hatching order. Instead, earlier-hatched chicks were more likely to outrun later-hatched siblings and reach the parent first.     

Effect of parentage and relatedness on the individual contribution to cooperative chick care in carrion crows Corvus corone corone

Carrion crows Corvus corone corone in northern Spain form complex cooperative groups that contain non-dispersing offspring and immigrants. Reproduction is often shared among group members, with polyandry prevalent over joint egg laying. However, due to incest avoidance or sexual immaturity, reproduction is potentially available only to a fraction of group members (potential breeders), while others do not breed (non-breeders). We combined molecular data with video-recorded observations at nests in order to investigate whether potential breeders and non-breeders adjust their individual effort in nestling feeding respectively to their level of parentage and the relatedness with the chicks. Overall, parents of at least one chick in the brood showed the highest feeding rates. Among potential breeders, genetic mothers and fathers fed chicks at significantly higher rates than individuals with no parentage, but they did not adjust their effort according to the proportion of offspring generated. Current direct fitness benefits are therefore important in determining a high provisioning effort, but crows may lack a mechanism to finely assess their share of parentage. Among non-breeders, males contributed more than females to chick feeding, but we found no significant correlation between feeding rate and relatedness to the nestling. We discuss how the latter result can be reconciled with the fact that kin selection has been shown to be important in shaping the crow cooperative society.Communicated by C. Brown     

Adaptations of meadow pipits to parasitism by the common cuckoo

Summary The meadow pipit Anthus pratensis is one of the most frequent hosts in Europe parasitized by the common cuckoo Cuculus canorus. The cuckoo normally removes one or more of the host eggs and replaces them with one of its own. The aim of the present study, which was conducted in an upland area of Central Norway, was to test the following question: assuming the cuckoo has laid a mimetic egg (which is slightly larger than and slightly different in color from that of a meadow pipit egg), under what circumstances are the parent meadow pipits able to detect such parasitism? The reaction of the meadow pipits to artificial parasitism was tested. Plastic model cuckoo eggs were used that bore a striking resemblance to real cuckoo eggs found in other meadow pipit nests in the same area. In addition, in some experiments a stuffed cuckoo dummy was used. The meadow pipits tolerated the experimental procedures and remained in their nests when given an artificial cuckoo egg, with or without removal of one of the host's eggs. However, when a host egg was removed and replaced with an artificial cuckoo egg, and at the same time a stuffed cuckoo dummy was presente, 50% of the birds deserted their nests. The difference between this result and the results of the other experiments was statistically significant. When only the stuffed cuckoo was presented, without any egg manipulations, the meadow pipits reacted in the same way as in the egg experiments. We conclude that the meadow pipit is capable of detecting whether or not its nest has been parasitized, provided it has observed the cuckoo near the nest site. Furthermore, because of the results of our experiments, we reject the hypothesis that the cuckoo has evolved egg removal behavior in order to prevent the host from assessing an increase in egg numbers.