A brood parasite selects for its own egg traits

Many brood parasitic birds lay eggs that mimic their hosts'' eggs in appearance. This typically arises from selection from discriminating hosts that reject eggs which differ from their own. However, selection on parasitic eggs may also arise from parasites themselves, because it should pay a laying parasitic female to detect and destroy another parasitic egg previously laid in the same host nest by a different female. In this study, I experimentally test the source of selection on greater honeyguide (Indicator indicator) egg size and shape, which is correlated with that of its several host species, all of which breed in dark holes. Its commonest host species did not discriminate against experimental eggs that differed from their own in size and shape, but laying female honeyguides preferentially punctured experimental eggs more than host or control eggs. This should improve offspring survival given that multiple parasitism by this species is common, and that honeyguide chicks kill all other nest occupants. Hence, selection on egg size in greater honeyguides parasitizing bee-eaters appears to be imposed not by host defences but by interference competition among parasites themselves.     

How selfish is a cowbird nestling?

Brood-parasitic young are reared in the nests of different species and can derive no genetic benefit from the survival of host offspring. However, although the nestlings of many parasitic cuckoo and honeyguide species routinely kill host young soon after hatching, nestling brown-headed cowbirds, Molothrus ater, tolerate host offspring and are commonly reared alongside them for at least part of the nestling period. I used comparative analyses of data from the literature to investigate whether brown-headed cowbird nestlings gain direct benefits by allowing host young to live. The brown-headed cowbird (44 g) parasitizes many passerines (adult mass range about 5-90 g) and the likelihood that host young survive to fledge from parasitized nests varies between species. In common with previous work, I found that host offspring mortality was highest in species whose offspring were relatively small compared with the cowbird nestling. Furthermore, cowbird nestlings were most likely to fledge when reared alongside host young of intermediate size. In these nests, one or two host young typically fledged as well. I suggest that cowbirds, and other host-tolerant brood parasites, could benefit from the presence of host nestlings through the assistance that host chicks offer in soliciting a higher provisioning rate, and that such benefits might outweigh the costs of having competition for food at each nest visit. Variation in this cost-benefit ratio could explain differences between brood parasite species in their tolerance of host young.     

Shiny Cowbird parasitism of a low quality host: effect of host traits on a parasite's reproductive success

ABSTRACT.   The reproductive success of parasitic cowbirds ( Molothrus spp.) varies among host species and is influenced by the degree of synchronization in timing of egg laying, the duration of parasite and host incubation periods, and the ability of hosts to incubate and rear parasite young. We studied the reproductive success of Shiny Cowbirds ( Molothrus bonariensis ) that parasitized the nests of Creamy-bellied Thrushes ( Turdus amaurochalinus ) in the Monte desert region of Argentina. Shiny Cowbirds frequently parasitized Creamy-bellied Thrush nests (60%), and most cowbirds synchronized egg laying with that of thrushes (79%). Most parasitic eggs (80%) hatched within 1 d of the hatching of the first host egg, and more than 91% of the eggs survived until the end of the incubation. However, only 60% of the cowbird eggs hatched and 52% of young survived. The proportion of Shiny Cowbirds eggs laid in Creamy-bellied Thrush nests that resulted in fledged young was 0.03, including eggs and young lost due to predation or desertion. Despite this low reproductive success, Creamy-bellied Thrushes were heavily parasitized by Shiny Cowbirds in our study area. Shiny Cowbirds may continue to parasitize these thrushes because of diffuse selection or because Shiny Cowbird chicks are more likely to fledge from Creamy-bellied Thrush nests in years or areas with greater food availability when brood reduction does not occur.     

Do shiny cowbird females adjust egg pecking behavior according to the level of competition their chicks face in host nests?

Interspecific brood parasites, like the shiny cowbird (Molothrus bonariensis), lay eggs in nests of other species. Shiny cowbird females peck and puncture eggs when they parasitize host nests. This behavior increases the survival of cowbird chicks when they have to compete for food with larger nestmates. However, cowbird chicks may benefit from smaller nestmates as they increase food provisioning by parents and the cowbird chicks secure most extra provisioning. We investigated whether egg-pecking behavior by female shiny cowbirds might be adjusted to the competition that their chicks face in host nests. We found that more host eggs are destroyed per cowbird egg laid in a larger-bodied host (chalk-browed mockingbird, Mimus saturninus, 70-75 g) than a smaller-bodied host (house wrens, Troglodytes aedon, 12-13 g). We also tested egg-pecking preferences in choice experiments with female cowbirds in captivity and found cowbirds presented with eggs in artificial nests pecked first and more frequently, and punctured more frequently the larger egg when this was a host egg, but not when this was a cowbird egg. Our results are partially consistent with the hypothesis that shiny cowbird females adaptively adjust their egg pecking behavior according to the competition that their chicks face in host nests.     

Cuckoo parasitism in a cavity nesting host: near absent egg‐rejection in a northern redstart population under heavy apparent (but low effective) brood parasitism

Brood parasite – host systems continue to offer insights into species coevolution. A notable system is the redstart Phoenicurus phoenicurus parasitized by the ‘redstart‐cuckoo’ Cuculus canorus gens. Redstarts are the only regular cuckoo hosts that breed in cavities, which challenges adult cuckoos in egg laying and cuckoo chicks in host eviction. We investigated parasitism in this system and found high overall parasitism rates (31.1% of 360 redstart nests), but also that only 33.1% of parasitism events (49 of 148 eggs) were successful in laying eggs into redstart nest cups. The majority of cuckoo eggs were mislaid and found on the rim of the nest; outside the nest cup. All available evidence suggests these eggs were not ejected by hosts. The effective parasitism rate was therefore only 12.8% of redstart nests. Redstarts responded to natural parasitism by deserting their nests in 13.0% of cases, compared to desertion rates of 2.8% for non‐parasitized nests. Our egg parasitism experiments found low rates (12.2%) of rejection of artificial non‐mimetic cuckoo eggs. Artificial mimetic and real cuckoo eggs added to nests were rejected at even lower rates, and were always rejected via desertion. Under natural conditions, only 21 cuckoo chicks fledged of 150 cuckoo eggs laid. Adding to this low success, is that cuckoo chicks are sometimes unable to evict all host young, and were more likely to die as a result compared to cuckoo chicks reared alone. This low success seems to be mainly due to the cavity nesting strategy of the redstart which is a challenging obstacle for the cuckoo. The redstart‐cuckoo system appears to be a fruitful model system and we suggest much more emphasis should be placed on frontline defences such as nest site selection strategies when investigating brood parasite–host coevolution.     

Function of egg punctures by Shiny Cowbirds in parasitized and nonparasitized Creamy‐bellied Thrush nests

ABSTRACT Avian brood parasites usually remove or puncture host eggs. Several hypotheses have been proposed to explain the function of these behaviors. Removing or puncturing host eggs may enhance the efficiency of incubation of cowbird eggs (incubation‐efficiency hypothesis) or reduce competition for food between cowbird and host chicks in parasitized nests (competition‐reduction hypothesis) and, in nonparasitized nests, may force hosts to renest and provide cowbirds with new opportunities for parasitism when nests are too advanced to be parasitized (nest‐predation hypothesis). Puncturing eggs may also allow cowbirds to assess the development of host eggs and use this information to decide whether to parasitize a nest (test‐incubation hypothesis). From 1999 to 2002, we tested these hypotheses using a population of Creamy‐bellied Thrushes (Turdus amaurochalinus) in Argentina that was heavily parasitized by Shiny Cowbirds (Molothrus bonariensis). We found that 56 of 94 Creamy‐bellied Thrush nests (60%) found during nest building or egg laying were parasitized by Shiny Cowbirds, and the mean number of cowbird eggs per parasitized nest was 1.6 ± 0.1 (N= 54 nests). At least one thrush egg was punctured in 71% (40/56) of parasitized nests, and 42% (16/38) of nonparasitized nests. We found that cowbird hatching success did not differ among nests where zero, one, or two thrush eggs were punctured and that the proportion of egg punctures associated with parasitism decreased as incubation progressed. Thus, our results do not support the incubation‐efficiency, nest‐predation, or test‐incubation hypotheses. However, the survival of cowbird chicks in our study was negatively associated with the number of thrush chicks. Thus, our results support the competition‐reduction hypothesis, with Shiny Cowbirds reducing competition between their young and host chicks by puncturing host eggs in parasitized nests.     

Nest Defense by Red Jungle Fowl (Gallus gallus spadiceus) Hens: The Roles of Renesting Potential,Parental Experience and Brood Reproductive Value

Reproductive-effort theory predicts that parents of any given age should expend more parental effort (1) as their residual reproductive value declines, and (2) as the reproductive value of offspring increases. An observational and experimental study of nest defense by captive red jungle fowl hens was used to examine these two predictions. Both young and old individuals significantly increased defense of the second nest compared to the first nest within a season; this pattern occurred for the defense of both eggs and chicks. Old hens showed significantly greater defense of both eggs and chicks in each of the nests than did young hens. Both young and old hens were significantly more defensive of chicks than eggs in each of two clutches of a season. Hens also reduced their nest defense significantly at the end of a two to three-day period after their chicks were replaced with eggs, and increased their nest defense after eggs were exchanged for chicks. Hens given four chicks showed more vigorous defense than hens given two chicks. When the brood size of hens with four chicks was reduced to one chick, the hens responded by exhibiting less vigorous nest defense. These patterns of nest defense in jungle fowl were not confounded by parental experience of hens, or differences in offspring quality that are related to time of breeding, maternal age, sire genetic quality or vulnerability of offspring to weather.     

Chick loss from mixed broods reflects severe nestmate competition between an evictor brood parasite and its hosts

Hatchlings of the obligate brood parasite common cuckoo Cuculus canorus typically evict eggs and nestmates but, rarely, host and parasite nestlings may grow up together. As part of previous experiments, we manipulated host clutches by inducing two great reed warbler Acrocephalus arundinaceus and one parasite young to share a nest from 4 days posthatch, when the cuckoo's eviction behaviour is thought to cease. We documented that in mixed broods typically at least one nestling eventually fell out of nest during the period of 5-10 days posthatch. In 83% of nests one or two host chicks disappeared, and in 17% of nests parasite chicks were lost. All nestlings remained in control broods of three hosts or one parasite. These results imply strong physical competition for space in mixed broods. We suggest that continued foster care for parasitized broods may occasionally be beneficial because host nestlings have some chance to escape the costs of parasitism, even when their parents fail to reject the parasite's egg and the parasite hatchling fails to evict nestmates. Conversely, evictor parasite chicks benefit not only through improved growth, as reported before, but also through the elimination of nestmate competition for space and the risk of displacement from mixed broods.     

Cuckoos versus reed warblers: Adaptations and counteradaptations

At study sites in Cambridgeshire, England, the percentage of reed warbler, Acrocephalus scirpaceus, nests parasitized by cuckoos, Cuculus canorus, in 2 years was 22·5% and 9·1%. The warblers rejected cuckoo eggs at 19% of parasitized nests. Parasitized clutches suffered less predation than unparasitized clutches, suggesting that the cuckoo itself was the major predator, plundering nests too advanced for parasitism so that the hosts would re-lay. The cuckoos laid a mimetic egg, parasitized nests in the afternoons during the host laying period, usually removed one host egg, laid a remarkably small egg and laid very quickly. Nests were experimentally parasitized with model eggs to study the significance of this procedure. Experiments showed that host discrimination selects for: (1) egg mimicry by cuckoos (poorer matching model eggs were more likely to be rejected); (2) parasitism during the laying period (mimetic eggs put in nests before host laying began were rejected); (3) afternoon laying (mimetic eggs were less likely to be accepted in the early morning than in the afternoon, when hosts were more often absent from the nest); (4) a small egg (large eggs, typical of non-parasitic cuckoos, were more likely to be rejected); (5) rapid laying (a stuffed cuckoo on the nest stimulated increased rejection of model eggs), and (6) sets a limit to host egg removal by cuckoos (if more than one or two are removed desertion may occur). Mimicry may also be selected for because it reduced the chance that second cuckoos can discriminate the first cuckoo's egg from the host's clutch. Predation did not select for mimicry; nests with a non-mimetic egg did not suffer greater predation than those with a mimetic egg. Host rejection of model eggs did not depend on: (1) stage of parasitism once host egg laying had begun (nevertheless cuckoos were more likely to lay early in the host laying period probably to increase the chance the cuckoo chick hatched); (2) removal of a host egg (however, this reduced the incidence of unhatched eggs so cuckoos may remove a host egg so as not to exceed the host incubation limit). There were two costs of rejection, an ‘ejection’ cost (own eggs ejected as well as the cuckoo egg) and, with mimetic eggs, a ‘recognition’ cost (own eggs ejected instead of the cuckoo egg). Reed warblers did not discriminate against unlike chicks (another species) and did not favour either a cuckoo chick or their own chicks when these were placed in two nests side by side. Possible reasons why the hosts discriminate against unlike eggs but not unlike chicks are discussed.     

Intraspecific nest parasitism in the Spotless Starling Sturnus unicolor

Intraspecific nest parasitism in two colonies of Spotless Starling Sturnus unicolor breeding in nestboxes was studied in central Spain from 1991 to 1994. Nests were monitored regularly and three criteria were used to detect nest parasitism: the appearance of more than one egg per day during the laying period of the host; the appearance of an egg after the start of incubation; eggs with unusual shape or pigmentation. The proportion of parasitized nests in first clutches (37%) was twice that of intermediate (19%) or second (20%) clutches in colony B, whereas parasitism occurred in first (35%) and intermediate (12%) but not in second clutches in colony A. Most clutches (52–70%) were parasitized during the host's laying period and received one parasitic egg. In 10% of the parasitized clutches in colony B, one of the host's eggs disappeared on the day the parasitic egg was added, suggesting that the parasitic female removed this egg. Although parasitism increased clutch size significantly, it led to a decrease in host breeding success, mainly through the removal of eggs and the loss of host nestlings and the survival of parasitic chicks. Observations suggested that parasitic females were young individuals without their own nests and/or those whose breeding attempt had been disrupted while laying in their own nest.     

Nestling discrimination without recognition: a possible defence mechanism for hosts towards cuckoo parasitism?

One of the great evolutionary puzzles is why hosts of parasitic birds discriminate finely against alien eggs, but almost never discriminate against parasitic chicks. A theoretical model has shown that an adaptive host response to alien eggs can be based on learning. However, learned nestling discrimination is too costly to be favoured by selection in hosts of evicting parasites, such as the European cuckoo (Cuculus canorus). Indeed, parasitic chick rejection has never been reported for any European cuckoo host species. As learned nestling discrimination is maladaptive, one can expect that a viable alternative for hosts would be to use discrimination mechanisms not involving learning and/or recognition. We suggest that hosts may starve and desert cuckoo chicks that require higher amounts of food than an average host brood at fledging (i.e. feeding rates to a parasite are outside the normal range of host behaviour in unparasitized nests). Our observations of the reed warbler (Acrocephalus scirpaceus) at parasitized nests indicate that such behaviour could possibly work in this host species.     

Partial host fidelity in nest selection by the shiny cowbird (Molothrus bonariensis), a highly generalist avian brood parasite

Obligate avian brood parasites can be host specialists or host generalists. In turn, individual females within generalist brood parasites may themselves be host specialists or generalists. The shiny cowbird Molothrus bonariensis is an extreme generalist, but little is known about individual female host fidelity. We examined variation in mitochondrial control region sequences from cowbird chicks found in nests of four common Argentinean hosts. Haplotype frequency distributions differed among cowbird chicks from nests of these hosts, primarily because eggs laid in nests of house wrens Troglodytes aedon differed genetically from those laid in nests of the other three hosts (chalk-browed mockingbird Mimus saturninus, brown-and-yellow marshbird Pseudoleistes virescens, and rufous-collared sparrow Zonotrichia capensis). These differences in a maternally inherited marker indicate the presence of a nonrandom laying behaviour in the females of this otherwise generalist brood parasite, which may be guided by choice for nest type, as house wrens nest in cavities whereas the other three species are open cup nesters.     

Intraspecific nest parasitism in the European starling Sturnus vulgaris

Biochemical genetic markers were used along with conventional methods (abnormal laying sequence/clutch size, unusual egg shape/pigmentation) to identify intraspecific nest parasitism at two British nestbox colonies of the European starling. Between 11 and 37% of first clutches were parasitized during 1977–1979. Parasitic females probably comprised all of the following categories: (1) paired females contesting a nestbox occupied by another pair; (2) previously paired females who had laid a clutch but had been unsuccessful; (3) unpaired females who had copulated with males that already had a mate and nest site; and (4) ‘professional’ nest parasites who distributed at lest some of their eggs in one or more nests other than their own. Although parasitized nests had higher clutch sizes, parasitism led to fewer host young fledging per egg laid, mainly through the eviction of eggs and subsequent nest desertion. Number of parasitic young fledged per egg laid was highest when eggs were laid synchronously with the host, when host clutches were larger, or a smaller number of parasite eggs were added to a nest, thus favouring parasites that distribute their eggs amongst a number of nests. A greater pressure on nest sites may have accounted for the higher levels of parasitism at the Aberdeen colony and for the greater number of parasite eggs laid in a nest. Although most parasitic female starlings appeared to be much less successful than non-parasitic ones, nest parasitism in the starling might evolve directly when one or more of the following advantages are present. (1) There are no constraints on the number of eggs a female may lay but there are constraints on the number of young she may feed adequately. (2) Female survival is increased by having fewer or no eggs/young to care for. (3) Current feeding conditions favour the survival of more young than would be produced by the most common clutch size. Intraspecific nest parasitism is considered to be a first stage in the evolution of interspecific nest parasitism.     

Competition with a host nestling for parental provisioning imposes recoverable costs on parasitic cuckoo chick's growth

Chicks of the brood parasitic common cuckoo (Cuculus canorus) typically monopolize host parental care by evicting all eggs and nestmates from the nest. To assess the benefits of parasitic eviction behaviour throughout the full nestling period, we generated mixed broods of one cuckoo and one great reed warbler (Acrocephalus arundinaceus) to study how hosts divide care between own and parasitic young. We also recorded parental provisioning behaviour at nests of singleton host nestlings or singleton cuckoo chicks. Host parents fed the three types of broods with similar-sized food items. The mass of the cuckoo chicks was significantly reduced in mixed broods relative to singleton cuckoos. Yet, after the host chick fledged from mixed broods, at about 10-12 days, cuckoo chicks in mixed broods grew faster and appeared to have compensated for the growth costs of prior cohabitation by fledging at similar weights and ages compared to singleton cuckoo chicks. These results are contrary to suggestions that chick competition in mixed broods of cuckoos and hosts causes an irrecoverable cost for the developing brood parasite. Flexibility in cuckoos' growth dynamics may provide a general benefit to ecological uncertainty regarding the realized successes, failures, and costs of nestmate eviction strategies of brood parasites.     

STUDIES OF THE DIEDERIK CUCKOO CHRYSOCOCCYX CAPRIUS IN THE TRANSVAAL

The migrant Diederik Cuckoo Chrysococcyx caprius was studied on a 72–acre area near the Klein Jukskei River near Johannesburg from 1955–1958. The birds first arrive in early October, after which the males take up territories and call continually until February; subsequent calls up to April are probably made by young birds.
Courtship displays, which involve courtship feeding, and egg-laying follow the birds' arrival by about a month. The Red Bishop Euplectes orix was the species most commonly parasitized, with smaller equal numbers of Cape Sparrows Passer melanurus and Masked Weavers Ploceus velatus . The cuckoos' eggs differ according to the host species and in two cases hatched between ten and 14 days after laying. The chicks normally evict their hosts' offspring on the second or third day after hatching. The fledging period appears to be roughly 19–20 days, and there is a period of post-fledging care lasting between 17 and 38 days, during which the chicks are fed different foods by different host species (grass seeds by bishops; insects of different sizes by weavers and sparrows).
The fact that the eggs and the calls of the chicks vary according to their host species suggest the existence of three separate host-specific strains in this area. The incidence of brood parasitism in Red Bishops' nests varied from 7–50% and averaged 25%.     

Breeding adaptations of Franklin's gull (larus pipixcan) to a marsh habitat

The behaviour and ecology of Franklin's gull were studied at Agassiz National Wildlife Refuge in northwestern Minnesota to determine the adaptations of the species for nesting in marshes. Two factors seemed to be important in colony site selection: cattail dispersion pattern and cattail density. Franklin's gulls prefer to nest in cattail areas closest to open water. The number of nests per unit area decreased as cattail density increased. Nest site selection is dependent on aggression and visibility. Visibility from nest level is the result of cattail placement and height. The distance between nests was directly correlated with visibility. Aggression by gulls on nests was lowered experimentally by decreasing visibility and raised by increasing visibility. Nest platforms were constructed of cattail material, and were attached to cattail stems. Nest material was added to the nests throughout the incubation and brooding period. Material was usually added following nest relief. The egg laying period was from 6 to 28 May. There was more synchrony of egg laying in sub-areas of the colony than in the colony as a whole. Successive eggs in clutches were laid at 24- to 48-hr intervals. The distance between nests decreased during the season as pairs filled in areas that were not defended. Territorial pairs defended an area up to 10 m from their stations prior to egg laying, but defended only the area within 3 m of their nests during incubation. Both members of pairs incubated the eggs and cared for the young. The incubation period was 24 days. The primary predators on adults and young were marsh hawk, great horned owl and mink. Franklin's gulls do not eat eggs or young of gulls. Adults fed on earthworms, insects and grain. Most marked adults fed within 16 km of the colony. Chicks were fed primarily on earthworms. The hatching period was from 30 May to 21 June. Chicks of all ages tested on a visual cliff apparatus were able to perceive the drop. Chicks tested on a 30-degree incline apparatus walked up it when 6 days old and younger, and walked down at 12 days of age and older. Brood mobility was less than in ground nesting species of gulls. In an undisturbed colony the chicks remained on the nest platforms until they were 25 to 30 days old although they were capable of swimming shortly after hatching. Individual recognition between parents and chicks appeared later in this species than in ground-nesting gulls. Adults accepted alien chicks (experimentally exchanged) that were younger than about 14 days old until their own chicks were over that age. Adults accepted larger and older broods than their own, as well as broods of mixed ages. Chicks began to react differently to strange adults at about 16 days of age. The breeding chronology of Franklin's gull is compressed when compared to that of other gulls. Possible selection pressures affecting this synchrony are discussed. The behaviour of the marsh-nesting Franklin's gull is compared with that of typical ground-and cliffnesting gulls; the possibility that the ancestral gull may have been a marsh nester is discussed.     

Behaviour of the yellow-eyed penguin chick

The ontogeny of yellow-eyed penguin ( Megadyptes antipodes ) chick behaviour follows the order of development determined by Nice (1962) for several species of birds, and by Spurr (1975) for the Adélie penguin ( Pygoscelis adeliae ). Feeding and comfort behaviours are the first to develop, followed by locomotion and aggressive behaviour.
Active solicitation of food may occur at one day of age. Chicks initially use non-visual cues to mediate begging. After their eyes open on the third or fourth day there is an increase in the use of visual stimuli, and begging occurs most often following adult nest relief. Sibling rivalry is not intense, occurring least during feeding, and in general both chicks are fed at each session.
The chicks are brooded for the first 21–25 days. At sparsely vegetated nest sites overheating may occur after 21 days and down-covered chicks will seek shade and pant in hot weather.
Throughout the 6–7 weeks of the guard phase there is a decrease in the amount of time spent resting in a prone posture, and an increase in exploratory, locomotory behaviour. During the post-guard phase, and until fledging and independence at 15 weeks after hatching, chicks may wander up to 20 m from the nest bowl during exploration, shade-seeking and feeding.
Adults feed only their own chicks, and chicks appear to beg only from their parents. Dense vegetation and long distances between nests tend to restrict contact between adults and chicks from neighbouring nests, and prevent the formation of large chick crèches.     

Brood parasite eggs enhance egg survivorship in a multiply parasitized host

Despite the costs to avian parents of rearing brood parasitic offspring, many species do not reject foreign eggs from their nests. We show that where multiple parasitism occurs, rejection itself can be costly, by increasing the risk of host egg loss during subsequent parasite attacks. Chalk-browed mockingbirds (Mimus saturninus) are heavily parasitized by shiny cowbirds (Molothrus bonariensis), which also puncture eggs in host nests. Mockingbirds struggle to prevent cowbirds puncturing and laying, but seldom remove cowbird eggs once laid. We filmed cowbird visits to nests with manipulated clutch compositions and found that mockingbird eggs were more likely to escape puncture the more cowbird eggs accompanied them in the clutch. A Monte Carlo simulation of this 'dilution effect', comparing virtual hosts that systematically either reject or accept parasite eggs, shows that acceptors enjoy higher egg survivorship than rejecters in host populations where multiple parasitism occurs. For mockingbirds or other hosts in which host nestlings fare well in parasitized broods, this benefit might be sufficient to offset the fitness cost of rearing parasite chicks, making egg acceptance evolutionarily stable. Thus, counterintuitively, high intensities of parasitism might decrease or even reverse selection pressure for host defence via egg rejection.     

The common cuckoo Cuculus canorus and its cavity nesting host, the redstart Phoenicurus phoenicurus: a peculiar cuckoo-host system?

We examined redstart Phoenicurus phoenicurus populations over a period of fifteen years to study interactions between the cuckoo Cuculus canorus and its cavity-nesting host. Over 380 redstart nests were checked and more than 100 cuckoo eggs were found during the study period. The average parasitism rate was 20%. The cuckoos' breeding success was extremely low, only 18 chicks surviving to the fledgling stage. When redstarts were parasitized experimentally with artificial cuckoo eggs, they rejected eight percent of mimetic eggs and 44% of non-mimetic eggs. We were not able to record any rejection of the real cuckoo eggs. However, about 30% of the real cuckoo eggs were found outside the redstart's nest cup. This could be the result of laying failures by the cuckoo, rather than of a strong rejection behaviour by the redstart. We suggest that redstarts' cavity nesting itself was a factor that reduced the cost of the parasitism dramatically. Firstly, it makes it difficult for the female cuckoo to lay her egg correctly in the nest and secondly, it is more difficult for the cuckoo chick to evict the host's eggs or nestlings effectively from the nest. Only 54% of the cuckoo chicks were able to evict all the host eggs or chicks from the nest. When reared in mixed broods, cuckoo chicks survived only in every second case to fledgling age, while at least one redstart chick from every brood managed to leave the nest.     

A review of the evidence for the translocation of eggs and young by nightjars (Caprimulgidae)

I undertook a review of evidence for the translocation of eggs and young by nightjars through a comprehensive search of the literature. I found that most of the evidence was based on hearsay, supposition, a misunderstanding of nightjar behaviour, or on the repetition of a story going back 200 years to Le Vaillant, via Audubon. There is no satisfactory direct evidence of any nightjar deliberately airlifting its eggs or young away from a disturbance. This conclusion is based on 10 studies by ornithologists across five continents, involving over 522 nests of 13 species. Accidental airlifting occurs occasionally when an egg or young chick gets stuck to the ventral plumage of the sitting adult. This is most likely to occur near the time of hatching. A nightjar can move an egg along the ground for a short distance, either by placing its lower mandible over the egg and then walking backwards while rolling it, or by rolling it forwards with its feet. Nightjars regularly move their chicks, especially after a disturbance. They do so by first moving away themselves and then calling. The chicks, who are highly mobile within hours of hatching, respond immediately by running to the parent birds.