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Egg retrieval is an ancient behaviour displayed by birds in which a displaced egg is transferred back into the nest. Egg rejection, in contrast, is a recently evolved adaptation by hosts of brood parasites in which an egg is removed from a nest. These conflicting behaviours are both adaptive because they prevent reproductive losses, but egg retrieval may benefit brood parasites if hosts retrieve parasitic eggs into their nests. We examined these conflicting cognitive processes in the prothonotary warbler (Protonotaria citrea), a cavity‐nesting host of the brown‐headed cowbird (Molothrus ater). Eggs and nestlings are occasionally found on the rims of warbler nest cavities. We determined how this happened using video cameras and tested whether warblers retrieved their own eggs more often than cowbird eggs. We also determined if shape affects retrieval and rejection by placing cylinders inside nest cups or onto nest rims. We found that eggs were displaced by rapidly departing warblers whose nests are constantly prospected by nest competitors and brood parasites. Warblers retrieved their eggs (29%) and cowbird eggs (19%) at similar rates, and never rejected cowbird eggs after retrieval. Warblers removed most cylinders from nest rims and cups (89% and 78%, respectively), and never retrieved them. Warblers have retained egg retrieval at a low level and its expression is costly because they only retrieve a low level of their own eggs and also retrieve cowbird eggs, which benefits the brood parasite. [ABSTRACT FROM AUTHOR]

Abandoning a brood to maximize lifetime reproductive success may result from the current costs affecting future reproductive opportunities. In certain contexts, clutch abandonment can be an evolved breeding strategy rather than a generalized response to stressors such as inclement weather and predation. Obligate brood parasitism, a reproductive strategy in which a parasitic species relies solely on other species to raise its young, imposes fitness costs to hosts and could serve as a trigger for clutch abandonment. This cost, and the resulting clutch abandonment strategy, may vary according to contact with the parasite, sensitivity to parasitism and the value of the current reproductive effort (i.e. brood value). We conducted a phylogenetically controlled meta-analysis covering 85 host species of the three generalist cowbird species (Molothrus sp.) in which we examined the following effects on the abandonment (nest desertion and egg burial) of parasitized versus unparasitized clutches: habitat at a small scale (higher host–parasite encounter frequency in open versus forested habitats) and at a large scale (longer interaction between parasites and hosts in prairie versus nonprairie regions), brood value (relative value of a clutch) and host species sensitivity to brood parasitism (relative body mass). Parasitism increased clutch abandonment overall. This increase was strongest in open nonforested habitats, with smaller, more sensitive hosts being more likely to abandon their clutch. Brood value and occurrence in prairie regions did not affect clutch abandonment, indicating that recent, more fine-scale host–parasite interactions were more important than coevolutionary history. Therefore, the abandonment of a brood can be used in diverse parental care strategies, including antiparasitic defences of brood-parasitic hosts. • We examined responses of hosts to parasitism by cowbirds (Molothrus sp.). • Parasitism increased clutch abandonment overall. • Abandonment strongly affected host sensitivity and current exposure to parasitism. • We found no effect of brood value or coevolutionary history. • Host behavioural plasticity may modulate host defences more than previously thought. [ABSTRACT FROM AUTHOR]

Egg retrieval is regarded as an ancestral behaviour that evolved in birds as an adaptation to eggs accidentally rolling out of nests and a fixed pattern of instinctive behaviour. By contrast, egg recognition is a later and specific adaptation that arises under selection pressure from brood parasitism, through which eggs likely belonging to brood parasites are rejected from the nest. Furthermore, egg recognition is flexible and can be enhanced through learning. We performed an empirical study on 12 populations of seven species in the tit family (Paridae), which revealed that mimetic eggs were retrieved more frequently than nonmimetic eggs across species; however, the rejection patterns were completely contrasting. Furthermore, species with a stronger recognition capacity of nonmimetic eggs retrieved eggs with lower probability; however, they rejected them with higher probability. This study also revealed that although egg retrieval is possibly a fixed pattern of instinctive behaviour, it can be modified to become a flexible reaction that adapts to subsequent evolution. Moreover, egg recognition, a specific adaptation to brood parasitism, gradually reshapes or modifies egg retrieval behaviour during evolution in a predictable manner, resulting in birds with higher levels of egg recognition being less willing to retrieve an egg from outside their nests. • Evolution of antiparasite behaviour reshapes egg retrieval, an ancestral behaviour. • Species good at egg recognition are less willing to retrieve eggs outside the nest. • Conspecific eggs are retrieved more frequently than foreign eggs across host species. • Foreign eggs are rejected more frequently than conspecific eggs across host species. [ABSTRACT FROM AUTHOR]

Hosts may evolve more than one egg type within a population (egg polymorphism) to make mimicry of any one individual more difficult. This process of co-evolution between cuckoos and their hosts has been well studied in certain species but has been ignored in other, lesser-known species. Using avian visual modelling, reflectance spectrophotometry, digital image analysis and egg volume, we examined the degree of egg mimicry between Klaas's cuckoo (Chrysococcyx klaas) and its host, the red-chested sunbird (Cinnyris erythrocercus), using museum collections. We found that the sunbird has evolved two distinct egg types based on luminance, colour and pattern, which we refer to as the light morph and the dark morph. Cuckoo eggs were significantly more similar to the dark morph eggs in luminance compared with the light morph eggs but were not more similar to dark morph eggs in colour, pattern or volume. To our knowledge, this is the first time that egg mimicry by Klaas's cuckoo has been studied objectively, and egg polymorphism was described in the red-chested sunbird host. Future studies focused on living populations would help us to gain a better understanding of the co-evolutionary interactions between this cuckoo and its host species. [ABSTRACT FROM AUTHOR]

What makes a perfect signature? Optimal signatures should be consistent within individuals and distinctive between individuals. In defense against avian brood parasitism, some host species have evolved "signatures" of identity on their eggs, comprising interindividual variation in color and pattern. Tawny-flanked prinia (Prinia subflava) egg signatures facilitate recognition and rejection of parasitic cuckoo finch (Anomalospiza imberbis) eggs. Here, we show that consistency and distinctiveness of patterns are negatively correlated in prinia eggs, perhaps because non-random, repeatable pattern generation mechanisms increase consistency but limit distinctiveness. We hypothesize that pattern properties which are repeatable within individuals but random between individuals ("invariant properties") allow hosts to circumvent this trade-off. To find invariant properties, we develop a method to quantify entire egg phenotypes from images taken from different perspectives. We find that marking scale (a fine-grained measure of size), but not marking orientation or position, is an invariant property in prinias. Hosts should therefore use differences in marking scale in egg recognition, but instead field experiments show that these differences do not predict rejection of conspecific eggs by prinias. Overall, we show that invariant properties allow consistency and distinctiveness to coexist, yet receiver behavior is not optimally tuned to make use of this information. [ABSTRACT FROM AUTHOR]

In response to brood parasitism, which significantly reduces the reproductive success of avian hosts, hosts have evolved the ability to recognize and reject parasitic eggs. Many studies have revealed the factors influencing egg recognition and egg rejection. Recent work has demonstrated that host personality along the bold–shy continuum can influence egg rejection behaviour. In addition, the egg rejection behaviour of hosts differs among populations, suggesting that individuals with different personalities might evolve various adaptive behaviours under different parasitic pressures, but there have been few relevant experiments to test this. In the present study, two populations of the Japanese tit, Parus minor , distributed in China were used to examine whether there were differences in egg rejection behaviour of individuals with different personalities. The results showed that egg rejection rates of bold individuals (which remained in the nest box when encountering human invaders) and shy individuals (which immediately flew away from the nest box when encountering human invaders) were similar in both populations of the Japanese tit, and there were also no significant differences in egg rejection rates between individuals of the same personality in the two populations. This study suggests that host personality along the bold–shy continuum does not account for egg rejection behaviour in Japanese tits. This might be attributable either to other factors that affect egg rejection behaviour counteracting the effect of personality or to the presence of multiple personality traits that act together in the Japanese tits, counteracting the effect of a single personality trait. [ABSTRACT FROM AUTHOR]

In host-parasite arms races, hosts can evolve signatures of identity to enhance the detection of parasite mimics. In theory, signatures are most effective when within-individual variation is low ('consistency'), and between-individual variation is high ('distinctiveness'). However, empirical support for positive covariation in signature consistency and distinctiveness across species is mixed. Here, we attempt to resolve this puzzle by partitioning distinctiveness according to how it is achieved: (i) greater variation within each trait, contributing to elevated 'absolute distinctiveness' or (ii) combining phenotypic traits in unpredictable combinations ('combinatorial distinctiveness'). We tested how consistency covaries with each type of distinctiveness by measuring variation in egg colour and pattern in two African bird families (Cisticolidae and Ploceidae) that experience mimetic brood parasitism. Contrary to predictions, parasitized species, but not unparasitized species, exhibited a negative relationship between consistency and combinatorial distinctiveness. Moreover, regardless of parasitism status, consistency was negatively correlated with absolute distinctiveness across species. Together, these results suggest that (i) selection from parasites acts on how traits combine rather than absolute variation in traits, (ii) consistency and distinctiveness are alternative rather than complementary elements of signatures and (iii) mechanistic constraints may explain the negative relationship between consistency and absolute distinctiveness across species.

Many hosts of brood-parasitic cuckoos reject foreign eggs from the nest. Yet where nests commonly receive more than one cuckoo egg, hosts might benefit by instead accepting parasite eggs. This is because cuckoos remove an egg from the nest before adding their own, and keeping cuckoo eggs in the nest reduces the odds that further host eggs are removed by subsequent cuckoos. This 'clutch dilution effect' has been proposed as a precondition for the evolution of cuckoo nestling eviction by hosts, but no previous studies have tested this in a host that rejects cuckoo nestlings. We tested the clutch dilution hypothesis in large-billed gerygones, Gerygone magnirostris , which are multiply parasitized by little bronze-cuckoos, Chalcites minutillus. Gerygones evict cuckoo nestlings but accept cuckoo eggs. Consistent with multiple parasitism favouring egg acceptance, we found gerygone egg survival was higher under scenarios of cuckoo egg acceptance than rejection. Yet gerygones were also flexible in their egg acceptance, with 61% abandoning clutches if they contained only cuckoo eggs. Our results thus confirm a clutch dilution effect in gerygones and suggest that multiple parasitism might favour a facultative response to cuckoo eggs, whereby hosts accept or reject cuckoo eggs depending on clutch composition. • The clutch dilution hypothesis was tested in large-billed gerygones. • We assessed whether retaining cuckoo eggs increases gerygone egg survivorship. • Gerygone egg survival was higher when hosts accepted cuckoo eggs. • Gerygones were also flexible in their response depending on clutch composition. • Multiply parasitized hosts tend to evolve both egg acceptance and nestling rejection. [ABSTRACT FROM AUTHOR]

Most mimicry systems involve imperfect mimicry, whereas perfect and high-fidelity mimicry are rare. When the fidelity of mimicry is high, mimics might be expected to have the upper hand against their antagonists. However, in coevolving systems, diversification of model phenotypes may provide an evolutionary escape, because mimics cannot simultaneously match all model individuals in the population. Here we investigate high-fidelity mimicry in a highly specialized, Afrotropical brood parasite–host system: the African cuckoo and fork-tailed drongo. Specifically, we test whether host egg polymorphisms are an effective defence against such mimicry. We show, using a combination of image analysis, field experiments and simulations, that: (1) egg colour and pattern mimicry of fork-tailed drongo eggs by African cuckoos is near-perfect on average; (2) drongos show fine-tuned rejection of foreign eggs, exploiting unpredictable pattern differences between parasitic eggs and their own; and (3) the high degree of interclutch variation (polymorphic egg 'signatures') exhibited by drongos gives them the upper hand in the arms race, with 93.7% of cuckoo eggs predicted to be rejected, despite cuckoos mimicking the full range of drongo egg phenotypes. These results demonstrate that model diversification is a highly effective defence against mimics, even when mimicry is highly accurate. [ABSTRACT FROM AUTHOR]

Interactions between avian brood parasites, such as common cuckoos (Cuculus canorus), and their hosts are one of the best‐studied examples of the coevolutionary arms race. Different stages of this arms race can be seen in different races of common cuckoos and their hosts across their range. However, little is known whether selected populations of two closely related but geographically distant species with probably different coevolutionary histories with the common cuckoo are also at different stages of the arms race. In this study, we tested this prediction experimentally using the same non‐mimetic model eggs and three‐dimensional (3D) printed models of the gray adult common cuckoo (Cuculus canorus). We examined egg recognition and egg rejection and aggression against the common cuckoo in the great reed warbler (Acrocephalus arundinaceus) and Oriental reed warbler (Acrocephalus orientalis), in Slovakia of Europe and northeast China of Asia. The results showed that the great reed warbler exhibited stronger responses to experimental model eggs and 3D models of the common cuckoo than the Oriental reed warbler. We conclude that both the great reed warbler and Oriental reed warbler have well‐developed antiparasitic behaviors against common cuckoos in the studied populations, but with different levels of defense intensity, which may be due to local differences in parasitic pressure and the risk of parasitism. This provides an opportunity to study coevolutionary processes between the brood parasite and its hosts together in both species at large geographical scales. [ABSTRACT FROM AUTHOR]

As a text-book example of coevolution, the escalating interactions between egg mimicry by parasitic cuckoos and egg recognition by their hosts constitute a key battlefield for parasitism and anti-parasitism strategies. However, some parasite–host systems have deviated from this coevolutionary trajectory because some cuckoos do not lay mimetic eggs, while the hosts do not recognize them, even under the high costs of parasitism. The cryptic egg hypothesis was proposed to explain this puzzle, but the evidence to date is mixed and the relationship between the two components of egg crypticity, egg darkness (dim egg coloration) and nest similarity (similarity to host nest appearance), remains unknown. Here, we developed a 'field psychophysics' experimental design to dissect these components while controlling for undesired confounding factors. Our results clearly show that both egg darkness and nest similarity of cryptic eggs affect recognition by hosts, and egg darkness plays a more influential role than nest similarity. This study provides unambiguous evidence to resolve the puzzle of absent mimicry and recognition in cuckoo–host systems and explains why some cuckoo eggs were more likely to evolve dim coloration rather than similarity to host eggs or host nests. [ABSTRACT FROM AUTHOR]

Our aim was to investigate the among-populations diversity of prey composition in Eurasian Reed Warblers' diets via their droppings, both to assess the ecological validity of this sampling method and to test whether the prey species most abundant in fecal samples were also the most present in the Italian study site. We collected fecal samples at four sites throughout Italy, for a total of 144 samples. Within reedbeds, the breeding habitat of the Eurasian Reed Warbler, we also collected arthropods by carrying out entomological sweepings at one of the study sites. Within the fecal samples, we identified dozens of prey species, belonging mainly to Araneae, Coleoptera, Diptera, Hymenoptera and Aphidoidea taxa, whose compositions were not statistically different among sites. The commonest prey species were Coleoptera in the droppings (68.5%) and Diptera in the reedbeds (31.3%), although the latter was less numerous in the fecal samples. The diets of different Italian populations of Eurasian Reed Warbler emerging from our study underline a strong Coleopteran presence, without differences across Italy. Fecal samples appear to be an exhaustive method to study variation in Eurasian Reed Warbler diet and its ecological importance; however, although potential caveats do exist, such as the possible under-representation of Diptera. [ABSTRACT FROM AUTHOR]

Simple Summary: Most birds build nests for laying eggs and rearing their offspring. They frequently clean their nests by removing foreign objects, such as leaves, small branches, stones, and feces. Further, some birds recognize and remove foreign eggs deposited by brood parasites. Nest sanitation and egg rejection are related behaviors as both impact brood survival and involve an accept/reject decision regarding the content of the nest. Here, we examined these behaviors in barn swallows (Hirundo rustica), one of the most abundant and widespread birds in the world. The results suggest that nest sanitation may be a more ancient behavior because it occurs more frequently, and foreign objects were removed sooner upon discovery than egg rejection. Therefore, nest sanitation may constitute an evolutionary precursor to foreign egg rejection; however, nest sanitation rarely increases egg rejection regarding probability and response latency. Female and male swallows engaged in nest sanitation and egg rejection, implying that both sexes are affected by this type of natural selection. Bird nests function as vessels for eggs and nestlings, and an environment for rearing offspring. However, foreign objects falling into bird nests and nestling eggshells may be harmful. Moreover, the smell of fecal sacs increases the risk of detection by predators. Many bird species have evolved nest sanitation to prevent damage to their nests. Furthermore, egg rejection evolved in some birds to thwart brood parasites that lay eggs in their nests. We studied 133 nests of barn swallows (Hirundo rustica) in an island population through a nest content manipulation experiment to determine nest sanitation and egg rejection behaviors and their relationship. Swallows rejected non-egg foreign objects more frequently (100% vs. 58.6%) and sooner than parasite eggs, which supports the hypothesis that nest sanitation is a pre-adaptation to egg rejection. However, nest sanitation did not increase egg rejection, either in probability or latency. Furthermore, both sexes incubated the eggs, cleaned the nests, and removed parasite eggs, implying that both are confronted with natural selection related to nest sanitation and brood parasitism. However, females invested more time in these behaviors than males. This provides evidence for the evolutionary relationship of nest sanitation and egg rejection behaviors in barn swallows. [ABSTRACT FROM AUTHOR]

Social learning can enable the rapid dissemination of behaviours throughout a population. Rejection of foreign eggs is a key defence used by hosts of avian brood parasites, and although learning has long been recognized as an important factor in egg rejection decisions, the role played by social cues remain less well understood. Here, we aimed to determine whether access to social information influenced egg rejection behaviours in semicolonial barn swallows, Hirundo rustica. By manipulating the social information available from a neighbouring nest, we showed that swallows with access to social information (i.e. a neighbour that recently rejected an egg) were more likely to reject a foreign egg that those that did not have access to social information (i.e. a neighbour that did not reject an egg). A subset of nests were videorecorded, and in these recordings the rejections were always performed by the incubating parent rather than a conspecific. This study provides direct evidence that egg rejection behaviours can be informed by social information in the absence of brood parasites, and in doing so highlights the dynamic nature of the defences that hosts deploy against brood parasitism. • Information about parasitism risk can inform host defences. • We tested whether social information could inform egg rejection decisions. • Barn swallows with access to social information rejected more foreign eggs. • Our results highlight the role that social information can play in coevolution. [ABSTRACT FROM AUTHOR]

Simple Summary: Obligate brood parasite birds such as the common cuckoo (Cuculus canorus) can trick their hosts via a variety of means. Cuckoos are threats to the host nests but not to the host adults themselves. To successfully parasitize the host nests, female cuckoos have been hypothesized to distract the hosts' attention from their nests by mimicking the calls of sparrowhawks (Accipiter nisus), a predator of the hosts. We performed playback experiments in two populations of the host oriental reed warbler (Acrocephalus orientalis) that have experienced different levels of parasitic pressure. We found that female cuckoo calls evoked both populations of the hosts to leave their nests more frequently than did the calls of male cuckoos or doves that do not pose threats to the hosts. This indicated that the call of the female cuckoo functions to deceive the host and thus favors host brood parasitism. However, we propose that such a deceiving effect of the female cuckoo call is due to the rapid cadence of the call rather than sparrowhawk mimicry. The common cuckoo (Cuculus canorus) is an obligate brood parasite that has evolved a series of strategies to trick its hosts. The female cuckoo has been hypothesized to mimic the appearance and sounds of several raptors to deceive the hosts into exhibiting anti-predator behavior. Such behavior would relax the protection of the host nest and thus allow the female cuckoo to approach the host nest unopposed. Many anti-parasite strategies have been found to vary among geographical populations due to different parasitic pressures from cuckoos. However, the effect of female cuckoo calls related to different levels of parasitic pressure has not been examined. Here, we studied the effect of female cuckoo calls on the oriental reed warbler (Acrocephalus orientalis), one of the major hosts of the common cuckoo, in two geographical populations experiencing different levels of parasite pressure. Four kinds of sounds were played back to the hosts: the calls from female common cuckoos, male common cuckoos, sparrowhawks (Accipiter nisus), and oriental turtle doves (Streptopelia orientalis). The results showed that the female cuckoo calls induced the hosts to leave their nests more frequently than the male cuckoo or dove calls in both populations, and two populations of the hosts reacted similarly to the female cuckoo calls, implying that the function of female cuckoo calls would not be affected by the difference in parasitism rate. This study indicates that female cuckoo calls function to distract the hosts' attention from protecting their nests. However, we propose that such a deception by the female cuckoo call may not be due to the mimicry of sparrowhawk calls, but rather that the rapid cadence of the call that causes a sense of anxiety in the hosts. [ABSTRACT FROM AUTHOR]

Visual complexity is ubiquitous in nature. Drivers of complexity include selection in coevolutionary arms races between antagonists. However, the causes and consequences of biological complexity and its perception are largely understudied, partly because complexity is difficult to quantify. Here, we address this by studying egg pattern complexity and its perception in hosts (tawny-flanked prinia Prinia subflava), which visually recognize and reject mimetic eggs of their virulent brood parasite (cuckoo finch Anomalospiza imberbis). Using field data and an optimization algorithm, we compute a complexity metric which predicts rejection of experimentally placed conspecific eggs in prinia nests. Real cuckoo finch eggs exhibit significantly lower pattern complexity than prinia eggs, suggesting that high complexity benefits hosts because it distinguishes host eggs from parasitic eggs. We show that prinias perceive complexity differences according to Weber's law of proportional processing (i.e. relative, rather than absolute, differences between stimuli are processed in discrimination, such that two eggs with simple patterns are more easily discriminable than two with complex patterns). This may influence coevolutionary trajectories of hosts and parasites. The new methods presented for quantifying complexity and its perception can help us to understand selection pressures driving the evolution of complexity and its consequences for species interactions. [ABSTRACT FROM AUTHOR]

Obligate avian brood parasites lay their eggs in the nest of other bird species, known as hosts. Brood parasitism often imposes severe fitness costs on hosts, selecting for the evolution of effective antiparasitic defences, such as recognition and rejection of brood parasite eggs. Glucocorticoids have been recently found to mediate host physiological and behavioral adjustments in response to brood parasite eggs; however, it remains unclear whether brood parasitism triggers a general response involving multiple physiological elements. In this study, we experimentally investigated whether a salient brood parasitic stimulus (the presence of a nonmimetic model egg in the nest) causes physiological adjustments in adult Eurasian blackbirds (Turdus merula) at immune (leukocyte profiles) and cellular (heat‐shock protein Hsp70 synthesis) level. Also, we explored whether these physiological changes are mediated by variations in corticosterone (CORT) levels. We found that experimental brood parasitism caused an increase in heterophils and a decrease in lymphocytes, leading to higher heterophils and lymphocytes ratios in parasitized birds. Nevertheless, we did not find tradeoffs between immune function and CORT levels. Hsp70 synthesis was not affected by our experimental manipulation. Our findings provide evidence that brood parasite eggs trigger a general stress response in egg‐rejecter hosts, including changes in cellular immune profiles. Highlights: Physiological effects of avian brood parasitism were assessed in adult hosts.Non‐mimetic brood parasite eggs led to increased heterophil/lymphocyte ratios, but not Hsp70 levels, in hosts.Changes on hosts' immune function were not mediated by plasma corticosterone. [ABSTRACT FROM AUTHOR]

Cuckoo nestlings thrive as avian brood parasites. To acquire sufficient food from the host parents, cuckoo nestlings generally make louder begging calls than host nestlings, but this may cause them to be more likely to attract the attention of predators. Studies have shown that nestlings would respond to the alarm calls of their parents by begging less, or crouching and remaining silent as an adaptation to reduce the risk of being heard by predators. Nevertheless, research is lacking on how parasite nestlings respond to alarm calls of their host parents. We studied the common cuckoo (Cuculus canorus) and one of the most common cuckoo host species, the oriental reed warbler (Acrocephalus orientalis), using a playback experiment in Yongnianwa National Wetland Park during the breeding seasons from June to July, 2020–2021. The begging behaviors of either cuckoo or host nestlings were quantified by playing back the alarm calls of host adults toward common cuckoo, sparrowhawk (Accipiter nisus), or oriental turtle doves (Streptopelia orientalis). Meanwhile, normal begging without playback, playback of the natural singing (NS) of host adults, and background noise (BN) were included as behavioral reference, non-threatening comparison, and playback control, respectively. The results showed that the cuckoo and host nestlings produced similar levels of begging with or without playback of NS and BN; however, both types of nestlings inhibited their begging intensity after hearing the playback of alarm calls, although they did not respond differently to the various alarm call playbacks. This study therefore elucidated that coevolution has selected the common cuckoo nestlings that adapt their begging behavior to the parent–offspring communication of alarm signaling in their host, oriental reed warblers.

Egg rejection in birds is a specific adaptation toward avian brood parasitism, whereas nest sanitation is a general behavior for cleaning the nest and avoiding predation. However, both behaviors refer to the action of ejecting objects out of the nest, and nest sanitation has been proposed as a pre-adaptation for egg rejection. Here, we tested the eliciting effect of nest sanitation on egg rejection in the red-whiskered bulbul Pycnonotus jocosus , a potential host species that are sympatric with parasitic cuckoos. We conducted meta-analyses of previous studies on both nest sanitation and egg rejection, in order to evaluate the consistency of our conclusions. Our results showed that nest sanitation did not elicit egg rejection in P. jocosus. The conclusions concerning such an eliciting effect from previous studies were mixed, whereas the methodologies were inconsistent, making the studies unsuitable for comparisons. However, the ejection frequency of nest sanitation was consistently higher than the frequency of egg rejection across different host species or populations. These results suggest that nest sanitation, which is an ancient behavior, is more fundamental than egg rejection, but the effect of the former on the latter is complex and needs further study. Standardized methodologies and the integration of behavior, physiology, and modeling may provide better opportunities to explore the relationship between nest sanitation and egg rejection. [ABSTRACT FROM AUTHOR]

Abstract Background Obligate brood parasites exert strong selective pressure on target hosts. In response, hosts typically evolve anti-parasitism strategies, of which egg recognition is one of the most efficient. Generally, host egg-recognition capacity is determined using model eggs. Previous studies have shown that some host species, which are capable of detecting parasite eggs, do not reject model eggs. However, it is unknown that whether the reaction to model eggs varies among distinct populations of the same host in relation to the degree of parasitism pressure. Results Here, we compared the rejection frequencies of model eggs and real eggs between mainland and island populations of the plain prinia (Prinia inornata), which are respectively sympatric and allopatric with their brood parasite, the common cuckoo (Cuculus canorus). Our results indicated that the mainland and island populations rejected real eggs at similar rates, but rejected model eggs, which were similar in size to real eggs but heavier, at significantly different rates: the island population rejected fewer model eggs, possibly because the rejection motivation of this population was lower due to absence of parasitism. Conclusions Our results indicated that some factors affecting the decision to reject, such as rejection motivation, varied according to the degree of parasitism pressure, and thus influenced the frequency of egg rejection. Furthermore, our results suggested that model eggs should be used with caution in comparative studies of egg recognition abilities among species or populations subjected to different intensities of brood parasitism. That is, model eggs may fail to accurately detect egg recognition in host populations with little to no risk of parasitism.

Parental care can be costly to parents’ fitness. As such, abandonment of the current reproductive attempt may benefit potential future opportunities, maximising lifetime reproductive success. Obligate brood parasitism, a reproductive strategy in which parasites lay their eggs in the nests of other species and rely solely on them to raise the parasitic young, is an ideal system to study brood abandonment. Some parasitised host species have evolved anti-parasitic defences, notably clutch abandonment (egg burial and nest desertion), that may mitigate negative consequences of parasitism. Abandonment of clutches due to parasitism is not unlike abandonment of reproduction in times of stress, suggesting that host responses to parasitism could be triggered at least partly by elevated stress hormones that mediate individual decisions. Yet, the mechanistic basis for clutch abandonment remains unclear. Here, we experimentally parasitised clutches of yellow warblers (Setophaga petechia), a common host of the brown-headed cowbird (Molothrus ater), with model cowbird eggs to examine whether host circulating corticosterone (CORT) differed among females that accepted parasitic eggs or rejected them through clutch abandonment. We also assessed whether feather CORT, a measure of past corticosterone exposure, differed between accepters and abandoners. Finally, we investigated whether egg visual signals, specifically differences in maculation characteristics between model cowbird and host eggs, predicted abandonment of experimentally parasitised clutches. Circulating CORT was higher in females who abandoned their parasitised clutches, but not in those who accepted, relative to controls with no egg addition. Past stress and differences in maculation characteristics did not predict whether individuals accepted or abandoned experimentally parasitised clutches. Moreover, differences in maculation characteristics between the host and model cowbird eggs did not predict CORT levels or nest abandonment. Thus, parasitism with subsequent clutch abandonment may be associated with elevated circulating CORT, but neither past stress nor differences in maculation characteristics influenced abandonment. The combination of these results contributes to our understanding of the roles of corticosterone and egg visual signals in the context of clutch abandonment in brood parasitism specifically, and of parental care more broadly.

Our aim was to investigate the among-populations diversity of prey composition in Eurasian Reed Warblers’ diets via their droppings, both to assess the ecological validity of this sampling method and to test whether the prey species most abundant in fecal samples were also the most present in the Italian study site. We collected fecal samples at four sites throughout Italy, for a total of 144 samples. Within reedbeds, the breeding habitat of the Eurasian Reed Warbler, we also collected arthropods by carrying out entomological sweepings at one of the study sites. Within the fecal samples, we identified dozens of prey species, belonging mainly to Araneae, Coleoptera, Diptera, Hymenoptera and Aphidoidea taxa, whose compositions were not statistically different among sites. The commonest prey species were Coleoptera in the droppings (68.5%) and Diptera in the reedbeds (31.3%), although the latter was less numerous in the fecal samples. The diets of different Italian populations of Eurasian Reed Warbler emerging from our study underline a strong Coleopteran presence, without differences across Italy. Fecal samples appear to be an exhaustive method to study variation in Eurasian Reed Warbler diet and its ecological importance; however, although potential caveats do exist, such as the possible under-representation of Diptera.

Bird nests function as vessels for eggs and nestlings, and an environment for rearing offspring. However, foreign objects falling into bird nests and nestling eggshells may be harmful. Moreover, the smell of fecal sacs increases the risk of detection by predators. Many bird species have evolved nest sanitation to prevent damage to their nests. Furthermore, egg rejection evolved in some birds to thwart brood parasites that lay eggs in their nests. We studied 133 nests of barn swallows (Hirundo rustica) in an island population through a nest content manipulation experiment to determine nest sanitation and egg rejection behaviors and their relationship. Swallows rejected non-egg foreign objects more frequently (100% vs. 58.6%) and sooner than parasite eggs, which supports the hypothesis that nest sanitation is a pre-adaptation to egg rejection. However, nest sanitation did not increase egg rejection, either in probability or latency. Furthermore, both sexes incubated the eggs, cleaned the nests, and removed parasite eggs, implying that both are confronted with natural selection related to nest sanitation and brood parasitism. However, females invested more time in these behaviors than males. This provides evidence for the evolutionary relationship of nest sanitation and egg rejection behaviors in barn swallows.

Hosts of brood parasitic cuckoos often employ mobbing attacks to defend their nests and, when mobbing is costly, hosts are predicted to adjust their mobbing to match parasitism risk. While evidence exists for fine-tuned plasticity, it remains unclear why mobbing does not track larger seasonal changes in parasitism risk. Here we test a possible explanation from parental investment theory: parents should defend their current brood more intensively as the opportunity to replace it declines (re-nesting potential), and therefore “counteract” any apparent seasonal decline to match parasitism risk. We take advantage of mobbing experiments conducted at two sites where reed warblers (Acrocephalus scirpaceus) experience (in Italy), or do not experience (in Finland), brood parasitism. We predicted that mobbing of cuckoos should be higher overall in Italy, but remain constant over the season as in other parasitised sites, whereas in Finland where cuckoos do not pose a local threat, we predicted that mobbing should be low at the beginning of the season but increase as re-nesting potential declined. However, while cuckoos were more likely to be mobbed in Italy, we found little evidence that mobbing changed over the season at either the parasitized or non-parasitized sites. This suggests that re-nesting potential has either little influence on mobbing behavior, or that its effects are obscured by other seasonal differences in ecology or experience of hosts.

In mimicry systems, receivers discriminate between the stimuli of models and mimics. Weber's Law of proportional processing states that receiver discrimination is based on proportional, not absolute, differences between stimuli. Weber's Law operates in a variety of taxa and modalities, yet it has largely been ignored in the context of mimicry, despite its potential relevance to whether receivers can discriminate models from mimics. Specifically, Weber's Law implies that for a given difference in stimulus magnitude between a model and mimic, as stimulus magnitudes increase, the mimic will be less discriminable from their model. This implies that mimics should benefit when stimulus magnitudes are high, and that high stimulus magnitudes will reduce selection for mimetic fidelity. Whether models benefit from high stimulus magnitudes depends on whether mimicry is honest or deceptive. We present four testable predictions about evolutionary trajectories of models and mimics based on this logic. We then provide a framework for testing whether receiver discrimination adheres to Weber's Law and illustrate it using coevolutionary examples and case studies from avian brood parasitism. We conclude that, when studying mimicry systems, researchers should consider whether receiver perception conforms to Weber's Law, because it could drive stimulus evolution in counterintuitive directions. [ABSTRACT FROM AUTHOR]

Abstract Background Our knowledge of avian brood parasitism is primarily based on studies of a few selected species. Recently, researchers have targeted a wider range of host–parasite systems, which has allowed further evaluation of hypotheses derived from well-known study systems but also disclosed adaptations that were previously unknown. Here we present developmental and behavioral data on the previously undescribed Plaintive Cuckoo (Cacomantis merulinus) nestling and one of its hosts, the Common Tailorbird (Orthotomus sutorius). Methods We discovered more than 80 Common Tailorbird nests within an area of 25 km2, and we recorded nestling characteristics, body mass, tarsus length and begging display every 3 days for both species. Results Plaintive Cuckoo nestlings followed a developmental pathway that was relatively similar to that of their well-studied relative, the Common Cuckoo (Cuculus canorus). Tailorbird foster siblings were evicted from the nest rim. The cuckoo nestlings gained weight faster than host nestlings, and required 3–9 days longer time to fledge than host nestlings. Predation was high during the early stages of development, but the nestlings acquired a warning display around 11 days in the nest, after which none of the studied cuckoo nestlings were depredated. The cuckoos’ begging display, which appeared more intense than that of host nestlings, was initially vocally similar with that of the host nestlings but began to diverge from the host sound output after day 9. Conclusions The developmental data on Plaintive Cuckoo nestlings and their tailorbird host builds an important foundation for future work on the co-evolutionary interactions in this parasite–host system.

The common cuckoo (Cuculus canorus) is an obligate brood parasite that has evolved a series of strategies to trick its hosts. The female cuckoo has been hypothesized to mimic the appearance and sounds of several raptors to deceive the hosts into exhibiting anti-predator behavior. Such behavior would relax the protection of the host nest and thus allow the female cuckoo to approach the host nest unopposed. Many anti-parasite strategies have been found to vary among geographical populations due to different parasitic pressures from cuckoos. However, the effect of female cuckoo calls related to different levels of parasitic pressure has not been examined. Here, we studied the effect of female cuckoo calls on the oriental reed warbler (Acrocephalus orientalis), one of the major hosts of the common cuckoo, in two geographical populations experiencing different levels of parasite pressure. Four kinds of sounds were played back to the hosts: the calls from female common cuckoos, male common cuckoos, sparrowhawks (Accipiter nisus), and oriental turtle doves (Streptopelia orientalis). The results showed that the female cuckoo calls induced the hosts to leave their nests more frequently than the male cuckoo or dove calls in both populations, and two populations of the hosts reacted similarly to the female cuckoo calls, implying that the function of female cuckoo calls would not be affected by the difference in parasitism rate. This study indicates that female cuckoo calls function to distract the hosts’ attention from protecting their nests. However, we propose that such a deception by the female cuckoo call may not be due to the mimicry of sparrowhawk calls, but rather that the rapid cadence of the call that causes a sense of anxiety in the hosts.

Theoretical studies predict that hosts of avian brood parasites should evolve defenses against parasitism in a matter of decades. However, opportunities to test these predictions are limited because brood parasites rarely switch to naïve hosts. Here, we capitalize on a recent host switch by the brood-parasitic Pacific Koel (Eudynamys orientalis) in eastern Australia, to investigate how quickly the Red Wattlebird (Anthochaera carunculata), a recent host that has been annexed by the koel within the last 90 years, can learn to recognize and mob adult cuckoos and evolve the ability to eject parasite eggs. Pacific Koel nestlings kill all host young, so there should be strong selection for hosts to evolve defenses. However, low parasitism rates and high egg recognition costs might slow the spread of egg ejection in our study populations, while adult parasite recognition should be able to spread more rapidly, as this defense has been shown to be a learned trait rather than a genetically inherited defense. We tested Red Wattlebirds at two sites where parasitism rate differed. As predicted, we found that the Red Wattlebird showed little or no ability to eject foreign model eggs at either site, whereas two historical hosts showed high levels of egg ejection at both sites. However, Red Wattlebirds responded significantly more aggressively to a koel mount than to mounts of a harmless control and nest predator at the site with the higher parasitism rate and gave significantly more alarm calls overall toward the koel mount. Our results support previous evidence that recognition and mobbing of a brood parasite are learned traits and may be especially beneficial to naïve hosts that have not had enough time or a high enough selection pressure to evolve egg rejection.

Although egg color polymorphism has evolved as an effective defensive adaptation to brood parasitism, spatial variations in egg color polymorphism remain poorly characterized. Here, we investigated egg polymorphism in 647 host species (68 families and 231 genera) parasitized by 41 species of Old Word cuckoos (1 family and 11 genera) across Asia, Europe, Africa, and Australia. The diversity of parasitic cuckoos differs among continents, reflecting the continent-specific intensities of parasitic selection pressure on hosts. Therefore, host egg polymorphism is expected to evolve more frequently on continents with higher cuckoo diversity. We identified egg polymorphism in 24.1% of all host species and 47.6% of all host families. The common cuckoo Cuculus canorus utilized 184 hosts (28.4% of all host species). Hosts of the common cuckoo and of Chrysococcyx species were more likely to have polymorphic eggs than hosts parasitized by other cuckoos. Both the number of host species and the host families targeted by the cuckoo species were positively correlated with the frequency of host egg polymorphism. Most host species and most hosts exhibiting egg color polymorphism were located in Asia and Africa. Host egg polymorphism was observed less frequently in Australia and Europe. Our results also suggested that egg polymorphism tends to occur more frequently in hosts that are utilized by several cuckoo species or by generalist cuckoo species. We suggest that selection pressure on hosts from a given continent increases proportionally to the number of cuckoo species, and that this selection pressure may, in turn, favor the evolution of host egg polymorphism. [ABSTRACT FROM AUTHOR]

Egg recognition is a variable but common anti‐parasitism defense among different species of birds with brood parasites. In contrast, nestling recognition is rare. Very few studies have found nestling recognition in brood parasite hosts and determined the rejection mechanism behind this behavior. Hosts may use the number of hatchling down‐feathers to reject parasite nestlings. We tested whether hatchling down‐feathers is a visual cue for the red‐rumped swallow, a host that can recognize and reject parasite nestlings. Our results indicated that red‐rumped swallows do not recognize foreign nestlings based on hatchling down‐feathers. The closed nest structure and hatchling morph may explain the absence of such a mechanism. None of the rejection mechanisms found in previous studies could explain the nestling recognition in swallows. Olfactory cues, tactile cues, or other visual cues, except for single nestling or hatchling down‐feathers, may provide nestling recognition in red‐rumped swallows. More study is needed to evaluate these possibilities. [ABSTRACT FROM AUTHOR]

Signal detection theory addresses the challenge of successfully identifying informative signals in noisy information contexts, allowing optimal behavioural decisions in diverse ecological contexts. The optimal acceptance threshold hypothesis proposed by Reeve (Reeve 1989 Am. Nat. 133, 407-435. (doi:10. 1086/284926)) is an elegant theoretical model to predict the flexibility of acceptance thresholds for conspecific discrimination. This model has provided a robust framework used to explore recognition systems in a broad range of contexts such as animal communication, nest-mate discrimination or anti-parasitic host responses. In this review, we discuss key concepts related to the optimal acceptance threshold hypothesis applied to egg rejection decisions in avian brood parasite-host interactions. We explore those factors determining signal detectability in parasitized nests and how hosts adjust their rejection decisions to both the risk of parasitism and the potential costs associated with egg rejection. In addition, we discuss recent results that challenge some traditional assumptions of the optimal acceptance threshold hypothesis and provide a novel perspective to explore rejection decisions, such as the existence of single-threshold decision rules or acceptance decisions. An integrative view combining current evidence with traditional theory is needed to further advance the comprehension of optimal acceptance thresholds. [ABSTRACT FROM AUTHOR]

We present a quantitative analysis ofthe data reported ic the only published review of parasitism frequency on hosts of Common Cuckoo cictüKî caaonir in a Medftenanean area We first eliminated a bias potentially introduced by the method by which data were recorded Of the minai potential "0 species parasitized in Italy, only 44 were confirmed as host species, of which only 10 species had more than 10% of their total nests parasitized We highlighted differential parasitism on host species according to geographic area, hot the analysis suggested results were strongly biased because nest Location was generally not reported and the number of records steeply decreased from North to South. [ABSTRACT FROM AUTHOR]

Background: Obligate brood parasites exert strong selective pressure on target hosts. In response, hosts typically evolve anti-parasitism strategies, of which egg recognition is one of the most efficient. Generally, host egg-recognition capacity is determined using model eggs. Previous studies have shown that some host species, which are capable of detecting parasite eggs, do not reject model eggs. However, it is unknown that whether the reaction to model eggs varies among distinct populations of the same host in relation to the degree of parasitism pressure. Results: Here, we compared the rejection frequencies of model eggs and real eggs between mainland and island populations of the plain prinia (Prinia inornata), which are respectively sympatric and allopatric with their brood parasite, the common cuckoo (Cuculus canorus). Our results indicated that the mainland and island populations rejected real eggs at similar rates, but rejected model eggs, which were similar in size to real eggs but heavier, at significantly different rates: the island population rejected fewer model eggs, possibly because the rejection motivation of this population was lower due to absence of parasitism. Conclusions: Our results indicated that some factors affecting the decision to reject, such as rejection motivation, varied according to the degree of parasitism pressure, and thus influenced the frequency of egg rejection. Furthermore, our results suggested that model eggs should be used with caution in comparative studies of egg recognition abilities among species or populations subjected to different intensities of brood parasitism. That is, model eggs may fail to accurately detect egg recognition in host populations with little to no risk of parasitism. [ABSTRACT FROM AUTHOR]

Cuckoos (Cuculus spp.) parasitize the nest of many host birds; however, it is not clear why some potential hosts that occupy the same area as cuckoos are rarely or not used as hosts. Potential hosts might adopt a variety of strategies to defend against cuckoo parasitism, such as attacking adult cuckoos to keep them away from the nest, rejecting cuckoo eggs, or recognizing and refusing to feed cuckoo nestlings. We investigated the strategies used by grey-backed thrushes (Turdus hortulorum), a potential host of cuckoos, to prevent cuckoo or conspecific brood parasitism. Our results indicated that the attack rate of grey-backed thrushes on cuckoo dummies was very low (11.4%). Thrushes showed egg polymorphism and exhibited 100% rejection of non-artificial blue model eggs. Thrushes also failed to identify the eggs (8.3% of rejection) of conspecifics. Thus, grey-backed thrushes show a high egg rejection towards nonmimetic eggs, but low towards conspecific eggs. We suggest that egg recognition and rejection in grey-backed thrushes may be a strategy to cope with interspecific brood parasitism rather than conspecific brood parasitism, and have probably evolved due to previous cuckoo parasitism. Significance statement: The explanations for high egg rejection in thrush (Turdus spp.) hosts were mixed without consistent conclusions. This study showed that grey-backed thrushes (Turdus hortulorum) show a high egg rejection towards nonmimetic eggs, but low towards conspecific eggs, suggesting that egg recognition and rejection in grey-backed thrushes may be a strategy to cope with interspecific brood parasitism rather than conspecific brood parasitism. [ABSTRACT FROM AUTHOR]

Avian brood parasitism is a model system for understanding coevolutionary arms races, and the great reed warbler ( Acrocephalus arundinaceus , hereafter 'warbler') and its parasite the common cuckoo ( Cuculus canorus , hereafter 'cuckoo') are prime examples of this coevolutionary struggle. Here, warblers select for egg colour mimicry by rejecting poorly matched cuckoo eggs. Contrary to long-held assumptions, recent work showed that warblers tend to reject lighter and browner eggs but tended to accept darker and bluer eggs rather than basing rejection decisions solely on perceived colour differences (i.e. the degree of mimicry). This counterintuitive, colour-biased rejection behaviour would select for bluer and darker cuckoo eggs, but would only be adaptive if cuckoos were consistently lighter and browner than warbler eggs. Therefore, we tested whether warbler eggs were consistently bluer and darker than cuckoo eggs. To do so, we re-analysed eggshell reflectance spectra of warblers and the cuckoos that parasitized them in the Czech Republic. As expected, we found that warbler eggs were significantly bluer and darker than the cuckoo eggs at the population level. Thus, we demonstrate imperfect mimicry in a long-coevolved cuckoo host-race and provide insights for exploring the coevolutionary interactions among hosts and their brood parasites.

Host defences against avian brood parasites are the outcome of welldocumented coevolutionary arms races, yet important questions about variation in hosts' antiparasitic response traits remain poorly understood. Why are certain defences employed by some species or individuals and not by others? Here, we propose that understanding variability in and the evolution of host defences can be facilitated by the study of the underlying physiological mechanisms. Specifically, because antiparasitic strategies involve behaviours that have been shown to be hormonally regulated in other contexts, we hypothesize that host responses to brood parasites are likely to be mediated by related endocrine mechanisms. We outline the hallmarks of the endocrine bases of parasite defence-related avian behaviours, review the current understanding of antiparasitic host tactics and propose testable hypotheses about the hormonal mechanisms that may mediate host defences. We consider these mechanisms in a life-history framework and discuss how endocrine factors may shape variation in host defences. By providing a hypothesis-driven mechanistic framework for defences against parasitism, this perspective should stimulate the study of their endocrine bases to enhance our understanding of the intricate arms races in avian host-parasite systems. [ABSTRACT FROM AUTHOR]

Abstract: Rothstein (Behavioral Ecology and Sociobiology, 11, 1982, 229) was one of the first comprehensive studies to examine how different egg features influence egg rejection behaviors of avian brood parasite–hosts. The methods and conclusions of Rothstein (1982) laid the foundation for subsequent experimental brood parasitism studies over the past thirty years, but its results have never been evaluated with replication. Here, we partially replicated Rothstein's (1982) experiments using parallel artificial model egg treatments to simulate cowbird (Molothrus ater) parasitism in American robin (Turdus migratorius) nests. We compared our data with those of Rothstein (1982) and confirmed most of its original findings: (1) robins reject model eggs that differ from the appearance of a natural robin egg toward that of a natural cowbird egg in background color, size, and maculation; (2) rejection responses were best predicted by model egg background color; and (3) model eggs differing by two or more features from natural robin eggs were more likely to be rejected than model eggs differing by one feature alone. In contrast with Rothstein's (1982) conclusion that American robin egg recognition is not specifically tuned toward rejection of brown‐headed cowbird eggs, we argue that our results and those of other recent studies of robin egg rejection suggest a discrimination bias toward rejection of cowbird eggs. Future work on egg recognition will benefit from utilizing a range of model eggs varying continuously in background color, maculation patterning, and size in combination with avian visual modeling, rather than using model eggs which vary only discretely. [ABSTRACT FROM AUTHOR]

In avian brood parasitism, both the host and the parasite are expected to develop various conflicting adaptations; hosts develop a defense against parasitism, such as an ability to recognize and reject parasitic eggs that look unlike their own, while parasites evolve egg mimicry to counter this host defense. Hosts may further evolve to generate various egg phenotypes that are not mimicked by parasites. Difference in egg phenotype critically affects the successful reproduction of hosts and parasites. Recent studies have shown that clear polymorphism in egg phenotype is observed in several host-parasite interactions, which suggests that egg polymorphism may be a more universal phenomenon than previously thought. We examined the mechanism for maintaining egg polymorphism in the rufescent prinia ( Prinia rufescens) that is parasitized by the plaintive cuckoo ( Cacomantis merulinus) from a theoretical viewpoint based on a mathematical model. The prinia has four distinct egg phenotypes: immaculate white, immaculate blue, white with spots, and blue with spots. Only two egg phenotypes, white with spots and blue with spots, are found in the cuckoo population. We show that the observed prinia and cuckoo phenotypes cannot be at an equilibrium and that egg polymorphism can be maintained either at stationary equilibrium or with dynamic, frequency oscillations, depending on the mutation rates of the background color and spottiness. Long-term monitoring of the prinia-cuckoo interaction over a wide geographic range is needed to test the results of the model analyses. [ABSTRACT FROM AUTHOR]

Recent studies have suggested that parasitic cuckoos have evolved laying behaviour resulting in matching of host and cuckoo eggs by choosing to lay eggs in host nests with host eggs that match the cuckoo eggs as an adaptation against egg recognition by the hosts. However, previous studies provided weak and indirect evidence with mixed results, leaving this question unresolved. Here, for the first time, we developed a robust methodology to provide unambiguous evidence that egg recognition in the host does not select for optimal egg matching during laying by the cuckoo. By using experiments that attracted parasitism, we showed that cuckoos, Cuculus canorus , indiscriminately laid eggs in oriental reed warbler, Acrocephalus orientalis , host nests containing real host eggs, egg-shaped models, stick models or coin models without any preference. Furthermore, cuckoos only selected to lay their eggs in nests with active hosts. These experiments provide evidence of cuckoos being indiscriminate in their choice of host nests, implying that coevolution of the egg phenotype of host and cuckoo eggs must have arisen from mechanisms other than matching of host eggs and those of the parasite. [ABSTRACT FROM AUTHOR]

The common cuckoo Cuculus canorus is a brood parasite that utilizes many host species. These have evolved defense against parasitism to reject cuckoo eggs that look unlike their own and some cuckoos have evolved egg mimicry to counter this defense. Egg phenotype indeed plays a key role for both the cuckoo and its hosts to successfully reproduce. It has been argued that cuckoos should parasitize host nests where egg phenotype matches because this makes parasitism more successful. Details of the cuckoo's parasitic behavior, however, largely remains unknown if they really parasitize hosts depending on 'egg matching'. In this paper, we model a time sequence of parasitic events in which a cuckoo finds host nests and decides to parasitize them or not in the presence of egg polymorphism. We evaluate which strategy is optimal: (1) opportunistic parasitism where cuckoos parasitize hosts irrespective of the phenotype, or (2) non-opportunistic parasitism where cuckoos parasitize hosts where egg phenotype matches. The analysis showed that either of the two strategies can be optimal. Factors not considered in the model, e.g., ecological and evolutionary changes both in the cuckoo and the host side, are discussed to explain apparent contrasts observed in some cuckoo-host interactions. [ABSTRACT FROM AUTHOR]

Brood-parasitic offspring sexually (mis)imprinting on the foster parents is considered one of the greatest constraints to the evolution of interspecific avian brood parasitism. While most nonparasitic juvenile birds learn the behaviours and mate choice preferences from their own parents, social parasites must avoid misimprinting on their host species' phenotype in order to accurately recognize conspecifics. One possible mechanism to assure accurate species recognition by juvenile parasites is to begin to associate with adult parasitic conspecifics, known as the ‘first contact’ scenario, whereby adult female parasites facilitate the dispersal of their offspring away from hosts, thus providing accurate referents for conspecific recognition. Using an automated radiotelemetry system, we determined the presence or absence (every 1–2 min during three breeding seasons; 516 315 search occasions) of radiotagged parasitic adult female brown-headed cowbirds, Molothrus ater , and compared their departures from a forest study site with those of genetic offspring or experimentally transplanted (nonrelated) juvenile cowbirds within the female's egg-laying range. Contrary to our predictions, we found no support for the facilitation of juvenile cowbird dispersal by adult female cowbirds. Juvenile cowbirds typically were not located within their mother's home range when departing the forest and, likewise, departure events for natal and experimentally transplanted juveniles (<2%) did not overlap temporally with the departure of the genetically assigned mother or with the departure of other local radiotagged female cowbirds. Surprisingly, we found that juvenile cowbirds primarily departed from the host's territory at sunset, when adult female cowbirds are infrequently present within the forest. Our results suggest that the solitary nocturnal roosting behaviour of juvenile cowbirds may facilitate independence from their hosts, thus minimizing the risk of misimprinting on heterospecific phenotypes. This strategy may also indirectly promote conspecific interactions, providing further evidence for the importance of independent spatial and social preferences of hosts and brood parasites in the evolution of avian brood parasitism. [ABSTRACT FROM AUTHOR]