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.     

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     

Shiny cowbirds share foster mothers but not true mothers in multiply parasitized mockingbird nests

Obligate brood parasitic birds, such as cowbirds, evade parental care duties by laying their eggs in the nests of other species. Cowbirds are assumed to avoid laying repeatedly in the same nest so as to prevent intrabrood competition between their offspring. However, because searching for host nests requires time and energy, laying more than one egg per nest might be favoured where hosts are large and can readily rear multiple parasites per brood. Such ‘repeat parasitism’ by females would have important consequences for parasite evolution because young parasites would then incur indirect fitness costs from behaving selfishly. We investigated shiny cowbird (Molothrus bonariensis) parasitism of a large host, the chalk-browed mockingbird (Mimus saturninus), in a population where over 70 % of the parasitized mockingbird nests receive multiple cowbird eggs. We assessed egg maternity directly, using cameras at nests to film the laying of individually-marked females. We also supplemented video data with evidence from egg morphology, after confirming that each female lays eggs of a consistent appearance. From 133 eggs laid, we found that less than 5 % were followed by the same female visiting the nest to lay again or to puncture eggs. Multiple eggs in mockingbird nests were instead the result of different females, with up to eight individuals parasitizing a single brood. Thus, while cowbird chicks regularly share mockingbird nests with conspecifics, these are unlikely to be their maternal siblings. Our results are consistent with shiny cowbird females following a one-egg-per-nest rule, even where hosts can rear multiple parasitic young.     

Post-ejection nest-desertion of common cuckoo hosts: a second defense mechanism or avoiding reduced reproductive success?

Hosts of the common cuckoo (Cuculus canorus), an avian brood parasite, develop antiparasite defense mechanisms to increase their reproductive success. Ejection of the parasite egg and desertion of the parasitized nest are the most typical adaptations in response to brood parasitism, but nest desertion may also occur in response to partial clutch reduction, independently from parasitism. Some great reed warblers (Acrocephalus arundinaceus) showed both mechanisms in the same incidence of cuckoo parasitism: in 18% of successful ejections of the parasite eggs, they deserted their nests. We studied if such cases of post-ejection nest-desertion are caused by brood parasitism or reduced clutch value. We experimentally parasitized clutches consisting of five or three host eggs with two painted conspecific eggs to mimic parasitic eggs, as multiple parasitism is frequent in the area. Although hosts ejected these parasitic eggs in both clutch categories (100% and 67% for the larger and smaller inital clutch sizes, respectively), we found that after manipulation, post-ejection nest-desertion frequently occurred at small (3-egg) clutches (40%), but rarely at large (5-egg) clutches (17%). The same phenomenon also occurred when unparasitized 3-egg clutches were reduced by two eggs, but not when 5-egg clutches were reduced in the same way. A logistic regression model revealed that only initial clutch size affected nest desertion of parasitized nests in our experiments. Therefore, we conclude that post-ejection nest-desertion is not a second antiparasite mechanism, which might serve as a redundant antiparasite defense, but a reaction to typically small and further decreased clutch size.     

Functional reference in an alarm signal given during nest defence: seet calls of yellow warblers denote brood-parasitic brown-headed cowbirds

Field observations and model-presentation experiments have shown that yellow warblers (Dendroica petechia) produce seet calls preferentially in response to brood-parasitic brown-headed cowbirds (Molothrus ater). In this study, we investigated whether seet calls are functionally referential alarm calls denoting cowbirds by determining whether female warblers responded appropriately to seet calls in the absence of a cowbird, whether alarm calling by warblers varied with response urgency, and how warblers in a population allopatric with cowbirds responded to cowbird and avian predator models and seet playbacks. As a control, we presented chip calls, which are elicited by nest predators as well as by non-threatening intruders, but are not strongly associated with cowbirds. Yellow warblers responded differently to playbacks of seet than chip calls. To seet playbacks, almost 60% of females gave seet calls and rushed to sit in their nests, responses typically elicited by cowbirds, whereas these responses were given infrequently in response to chip calls. Yellow warblers seet called equally in situations that simulated low, medium and high risk of parasitism, which suggests that these calls did not vary with response urgency. In a population allopatric with cowbirds, seet calls were rarely produced in response to cowbird or avian nest predator models and never to seet playbacks. These results suggest that seet calls are functionally referential signals denoting cowbirds and that cowbird parasitism was a strong selective pressure in the evolution of functional referentiality in the seet call of yellow warblers.Communicated by W.A. Searcy     

Age,experience, and enemy recognition by wild song sparrows

Summary Older female Song Sparrows (Melospiza melodia) breeding on Mandarte Island, B.C., Canada, are more often parasitized by a brood parasite, the Brown-headed Cowbird (Molothrus ater), than yearling females breeding for the first time. This may be explained if older Song Sparrows behave differently than yearlings towards searching female cowbirds, and are thus more readily recognized as potential hosts. We tested this hypothesis by measuring the responses of Song Sparrows to a stuffed mount of a female cowbird in 1982, when no cowbirds frequented the island. This mount, and a control (a female junco) were presented near the nests of wild female sparrows of known age. As predicted, adult female sparrows behaved differently towards the cowbird model from yearlings (Table 1). They did not differ in their responses to the junco. Adult males also responded differently to the cowbird and junco (Table 2), but adult males did not differ significantly from yearlings in their response to either model. When birds that responded weakly as yearlings in the absence of cowbirds were retested as 2-year-olds after the recolonization of the island by cowbirds, they responded strongly to cowbird models. Two-year-old birds not exposed to cowbirds as yearlings, were parasitized at a rate intermediate between the rate for experienced adults and that for yearlings. Yearling females were parasitized less often at the beginning of the period of breeding activity by cowbirds than at the end. All these results are consistent with our hypothesis that age-selective parasitism results from differences among age classes in the mobbing responses of Song Sparrows to cowbirds. The greater response of adult sparrows to cowbirds seems non-adaptive, because it apparently results in a loss of reproductive output through selective parasitism. We suggest that this paradox can be resolved if the response to cowbirds near the nest is an instance of a more general acquired response to all potential enemies that approach a nest.     

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.     

Costs of brood parasitism and the lack of defenses on the yellow-winged blackbird - shiny cowbird system

The shiny cowbird (Molothrus bonariensis) is a generalist brood parasite that lays either white-immaculate or spotted egg morphs in eastern Argentina and Uruguay. Some hosts accept both morphs, others accept spotted eggs and reject the white morph, but no host has been found to accept white eggs and reject spotted ones. It has been suggested that the yellow-winged blackbird (Agelaius thilius) may be that type of host. The finding of a white acceptor-spotted rejector species would help to explain the occurrence and maintenance of the parasite egg polymorphism. We studied the incidence of shiny cowbird parasitism on this host, its costs for their reproductive success and the presence of antiparasitic defenses in the yellow-winged blackbird - shiny cowbird system. The parasite affected the reproductive success of the host in two ways. Cowbirds punctured host eggs causing a reduction in clutch size, and yellow-winged blackbirds deserted their nests whenever they suffered high egg loss. In addition, parasitized nests suffered higher predation during the nestling stage, but not during egg stages, indicating that the difference found was related to the presence of the cowbird chick, and not to higher exposure of parasitized nests to both parasites␣and predators. Despite the costs imposed by the parasite, yellow-winged blackbirds have not evolved antiparasitic defenses. This host did not reject any egg morph of the shiny cowbird nor desert parasitized nests unless it had suffered high egg loss. Current explanations for the host lack of defenses, the “time lag” and the “equilibrium” hypothesis, are discussed. Received: 29 August 1997 / Accepted after revision: 10 January 1998     

Cowbird removals unexpectedly increase productivity of a brood parasite and the songbird host

Generalist brood parasites reduce productivity and population growth of avian hosts and have been implicated in population declines of several songbirds of conservation concern. To estimate the demographic effects of brood parasitism on Bell's Vireos (Vireo bellii), we removed Brown-headed Cowbirds (Molothrus ater) in a replicated switchback experimental design. Cowbird removals decreased parasitism frequency from 77% and 85% at unmanipulated plots to 58% and 47% at removal plots in 2004 and 2005, respectively. Vireo productivity per pair was higher at cowbird removal plots when years were pooled (mean = 2.6 +/- 0.2 [SE] young per pair) compared to unmanipulated plots (1.2 +/- 0.1). Nest desertion frequency was lower at cowbird removal plots (35% of parasitized nests) compared to unmanipulated plots (69%) because removal of host eggs was the proximate cue for nest desertion, and vireos experienced lower rates of egg loss at cowbird removal plots. Nest success was higher among unparasitized than parasitized nests, and parasitized nests at cowbird removal plots had a higher probability of success than parasitized nests at unmanipulated plots. Unexpectedly, cowbird productivity from vireo pairs was higher at cowbird removal plots (mean = 0.3 +/- 0.06 young per pair) than at unmanipulated plots (0.1 +/- 0.03) because fewer parasitized nests were deserted and the probability of nest success was higher. Our study provides the first evidence that increases in cowbird productivity may be an unintended consequence of cowbird control programs, especially during the initial years of trapping when parasitism may only be moderately reduced. Thus, understanding the demographic impacts of cowbird removals requires an informed understanding of the behavioral ecology of host-parasite interactions.     

Cues used by shiny cowbirds (Molothrus bonariensis) to locate and parasitise chalk-browed mockingbird (Mimus saturninus) nests

Unlike other birds, shiny cowbirds (Molothrus bonariensis) must locate host nests where to lay their eggs and then decide whether to parasitise them. They should also synchronise their laying with that of the host to increase the survival of parasite egg and young. Shiny cowbirds can discover nests using host behaviour as a cue, or by searching the habitat without need for the presence of a host. Besides, they can synchronise parasitism with host laying by monitoring nests during building and laying, or directly by assessing the degree of development of embryos through the puncture of host eggs. Alternatively, synchronization can arise by lower nest attentiveness during host laying. We determined the extent of synchronization between laying of shiny cowbirds and chalk-browed mockingbirds (Mimus saturninus) and estimated if parasitism was negatively associated with host nest attentiveness. We also conducted an experiment to test if host activity was necessary to locate nests, and if puncture of host eggs was a cue for deciding parasitism. Shiny cowbirds synchronised parasitism with host laying in 75% of the cases and synchronization was not explained by lower host nest attentiveness during laying. Shiny cowbirds located nests without need for presence of a host, but the decision of parasitising the nest depended on host activity at the nest. The information that shiny cowbirds could obtain through egg punctures was not necessary for deciding parasitism. Our results indicate that shiny cowbirds rely on the precise timing of their eggs and avoid laying in unsuitable nests.     

Local Gradients of Cowbird Abundance and Parasitism Relative to Livestock Grazing in a Western Landscape

Abstract: We studied local patterns of Brown-headed Cowbird ( Molothrus ater ) abundance, parasitism rates, and nest success of a common host, the Plumbeous Vireo ( Vireo plumbeus ), in relation to the distribution of livestock grazing in an undeveloped region of northeastern New Mexico, 1992–1997. We predicted that both cowbird abundance and parasitism rates of vireo nests would decrease with increasing distance from active livestock grazing, and that the nesting success of vireos would increase. We measured cowbird abundance and host density and located and monitored vireo nests in pinyon-juniper and mixed-conifer habitats that ranged from actively grazed to isolated from livestock grazing by up to 12 km. Cowbird abundance declined with distance from active livestock grazing and was not related to host density or habitat type. Brood parasitism levels of vireo nests ( n = 182) decreased from> 80% in actively grazed habitats to 33% in habitats that were 8–12 km from active grazing but did not vary by habitat type or distance to forest edge. Vireo nesting success was higher in mixed-conifer habitat than in pinyon-juniper but was unrelated to distance from active livestock grazing. Nest losses due to parasitism declined with distance from active livestock grazing. Our results suggest that cowbird abundance and parasitism rates of hosts may be distributed as a declining gradient based on distance from cowbird feeding sites and that isolation from feeding sites can reduce the effects of parasitism on host populations. These findings provide support for management techniques that propose to reduce local cowbird numbers and parasitism levels by manipulating the distribution of cowbird feeding sites. The presence of parasitized nests> 8 km from active livestock grazing suggests that, in some regions, management efforts may need to occur at larger scales than previously realized.     

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     

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.     

Is reduced clutch size a cost of parental care in Eastern Phoebes (Sayornis phoebe)?

What is the cost of parental care in birds? Previous studies using observational and experimental techniques on nest building and clutch sizes in a small migrant flycatcher, the Eastern Phoebe (Sayornis phoebe), led to contradictory results that did not show a consistent cost of current reproductive effort on residual reproductive output. The data presented here indicate that different elements of parental behaviors are indeed costly because they reduce various aspects of phoebes' subsequent reproductive performance. Experimental removal of old nesting structures at previously used breeding sites reduced but did not eliminate the chance of phoebes' settlement in the subsequent year. Comparing sites at which phoebes did and did not build new nests showed that nest builders completed their first clutches later, had lower probabilities of second breeding attempts, and more often lost their nesting attempt due to fallen nest structures than nest reusers. There was, however, no significant effect of nest building on the clutch sizes and rates of cowbird parasitism of first nesting attempts. Overall, sites with newly built nests had lower seasonal reproductive effort than sites with reused nests. I also examined phoebes' relative residual reproductive output in a separate breeding season when nest building was not experimentally manipulated. When controlled for confounding variables this analysis indicated that in those phoebes that did breed for a second time, the relative decrease of the sizes of first to presumed second clutches was greater at sites where first breeding attempts consisted of more total nestlings. These data are consistent with the hypothesis that parental care is costly in Eastern Phoebes and support predictions of trade-offs between the nest building, brood care, and residual egg-investment components of reproduction.     

Parasitism at the Landscape Scale: Cowbirds Prefer Forests

Landscape-scale examination of parasitism patterns of Brown-headed Cowbirds ( Molothrus ater ) revealed heterogeneous parasitism rates across the mosaic of a forest and associated old-field communities. In a two year study in Dutchess County, New York, we found a significantly higher parasitism rate in the forest-interior community ( n = 301 nests; 17 species) than on the species in the adjacent and nearby old field and edge ( n = 328 nests; 15 species; 32.3% versus 6.5%; p < 0.0001). Cowbirds invaded a mature 1300-ha forest stand even when their traditional host species were available in adjacent old-field and edge habitats. The forest and old-field study areas were located in a 38,000-ha township with 55% forest cover and contained numerous agriculture, dairy, and horse farms that provided favorable habitat for cowbirds. Within-forest examination of parasitism patterns revealed four aspects of cowbird parasitism that contrasted with patterns described in other regions: (1) parasitism was concentrated significantly more often on ground- and low-nesting (nests ≤ 1 m) forest species than on medium- and high-nesting species (nests> 1 m; 35.01% versus 29.93%; p = 0.0393); (2) parasitism was not significantly greater on Neotropical migrant species than on short-distance migrants and residents; (3) the parasitism rate was not higher in nests close to edges; and (4) the parasitism level was low on certain forest species (such as Wood Thrush) that have experienced high parasitism levels in the Midwest. From a management perspective these data suggest that cowbirds exhibit regional differences in host and habitat use; the target host community of a particular cowbird population is unpredictable at the landscape scale; and a landscape scale should be used in designing cowbird studies to accurately assess local population dynamics.     

Mechanism of egg recognition in defenses against conspecific brood parasitism: American coots (<Emphasis Type="Italic">Fulica americana</Emphasis>) know their own eggs

Hosts of avian brood parasites use a variety of defenses based on egg recognition to reduce the costs of parasitism; the most important of which is rejecting the parasitic eggs. Two basic recognition mechanisms are possible: “true recognition”, whereby hosts recognize their own eggs irrespective of their relative frequency in the clutch, and minority recognition (or “recognition by discordancy”), whereby hosts respond to the minority egg type. The mechanism of recognition has been experimentally studied in a handful of species parasitized by interspecific brood parasites, but the mechanism used in defenses against conspecific brood parasitism is unknown. I experimentally determined the mechanism of egg recognition in American coots (Fulica americana), a species with high levels of conspecific brood parasitism, egg recognition, and rejection. I swapped eggs between pairs of nests to alter frequencies of host and “parasite” eggs and then used two criteria for recognition: egg rejection and nonrandom incubation positions in the clutch. Eight of 12 nests (66%) given equal frequencies of host and parasite eggs showed evidence of true recognition. In contrast, only one of eight (12.5%) nests where host eggs were in the minority showed evidence of recognition by discordancy. The nonrandom incubation positions of parasitic eggs indicates that birds sometimes recognize parasitic eggs without rejecting them and provides a means of assessing recognition on a per nest basis in species with large clutches. Adaptive recognition without rejection may also be an important evolutionary stepping stone to the evolution of egg rejection in some taxa.     

Utilization of a new host in the screaming cowbird <Emphasis Type="Italic">Molothrus rufoaxillaris</Emphasis>, a host specialist brood parasite: host switch or host acquisition?

The screaming cowbird Molothrus rufoaxillaris has been long known as a host specialist brood parasite. However, in the past years, the utilization of two new hosts has been documented. We examined the variation in mitochondrial control region sequences from screaming cowbird chicks found in the nests of two hosts, the bay-winged cowbird (Agelaioides badius), which is its regular host, and the chopi blackbird (Gnorimopsar chopi), which is a new host, in Formosa Province, Argentina. If a group of females switched to this new host, we expected to find an association between host use and haplotype frequency distribution, indicating the presence of host-specific female lineages, whereas we expected no such association if the cowbird population incorporated this new host and females use both hosts simultaneously. Haplotype frequency distributions differed between cowbird chicks from the nests of both hosts. This indicates that nest choice by females of this brood parasite is not random and that they preferentially parasitize the nests of the same host species.     

Foreign egg retention by avian hosts in repeated brood parasitism: why do rejecters accept?

Great reed warblers (Acrocephalus arundinaceus) are frequently parasitized by egg-mimetic common cuckoos (Cuculus canorus) in Hungary, and these hosts reject about a third of parasitic eggs. The timing of parasitism is important, in that the probability of rejection decreases with advancing breeding stages in this host. Also, egg rejection is more common when a clutch is parasitized by a single foreign egg, compared to parasitism by multiple eggs. We repeatedly parasitized great reed warbler clutches with moderately mimetic foreign eggs, either with (1) one foreign egg (single parasitism) and, after 3 days, by all foreign eggs (multiple parasitism), or (2) all foreign eggs and, 3 days later, by only one foreign egg. Hosts ejected 26–53 % of the experimental parasitic eggs in the first stage of the repeated parasitism, but almost all eggs were accepted in the second stage, irrespective of whether the clutch was singly or multiply parasitized. Video-taping of the behavioural responses of hosts to experimental parasitism revealed no evidence for sensory constraints on foreign-egg recognition, because hosts recognized and pecked the parasitic eggs as frequently in the second stage of repeated parasitism, as they did in the first stage. We suggest that the relative timing of parasitism (laying vs. incubation stage), rather than learning to accept earlier-laid foreign eggs, results in higher acceptance rates of cuckoo eggs in repeated parasitism, because there is decreasing natural cuckoo parasitism on this host species and, hence, less need for antiparasitic defences, with the advancing stages of breeding.     

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.     

Egg adoption can explain joint egg-laying in common eiders

Hypotheses regarding the evolution and maintenance of intraspecific nest parasitism were tested with data collected during a 3-year study of common eiders (Somateria mollissima) breeding near Churchill, Manitoba. The nest parasitism rate was highest (42.4% of nests) during the year with the highest nest density and the best environmental conditions, and lowest (20.2% of nests) in the year with the lowest nest density and the poorest environmental conditions. Over the nesting season, parasitic eggs were laid at the same time as normally laid eggs. Most parasitic eggs (>75%) were laid before the host female laid her third egg. The majority of the parasitic eggs were the first or second egg produced by the parasitic female. When a parasitic egg was laid before or on the same day as the host female initiated her clutch, the probability of her first egg being depredated before incubation was significantly lowered. First- and second-laid eggs suffered a high rate of predation probably because nesting females do not attend their clutch until their second or third egg is laid. Hypotheses that some females use intraspecific nest parasitism to parasitize the parental care of other females were inconsistent with these data. Egg adoption is a likely explanation for the prevalence of females incubating parasitic eggs in this population. Received: 30 September 1997 / Accepted after revision: 6 May 1998