In predator-prey interactions, the prey faces extreme challenges from predation, which drives the evolution of defense or anti-predator mechanisms. Compared with adult birds, nestlings are more vulnerable but not helpless. However, data on whether nestlings eavesdrop on the danger signals transmitted by other prey nestlings and the mechanisms of eavesdropping remain limited. In brood parasitism, common cuckoo (Cuculus canorus) nestlings, raised by host adults who are not closely related, offer an instructive system for studying the transmission and recognition of danger signals among nestlings of different species that share special relationships. We played back the distress calls of common cuckoo nestlings to nestlings of three sympatric host species (the oriental reed warbler Acrocephalus orientalis, which is a primary host of the common cuckoo, the reed parrotbill Paradoxornis heudei, an occasional host, and the vinous-throated parrotbill Sinosuthora webbiana, which is not parasitized in the study area) to investigate whether the host nestlings reduced their begging behavior. We also quantified the degree of inhibition toward begging behavior for these nestlings. The results revealed that, in response to the distress calls, the three sympatric host species markedly suppressed their begging behavior. This response can likely be attributed to the innate response of host nestlings caused by the general characteristics of distress calls, rather than the acoustic similarity and phylogenetic relationship between host nestlings and cuckoo nestlings. Furthermore, we observed that upon hearing the distress calls of cuckoo nestlings, the oriental reed warbler nestlings exhibited the greatest reduction in the total number of calls compared to the other two host species, potentially owing to stronger predation and parasitic pressures. This study suggests that host nestlings can detect danger signals emitted by parasitic nestlings; however, further investigation is needed to determine whether they can respond to distress calls from unfamiliar nestlings in different regions.

When host nests are scarce, avian brood parasites would benefit from behaviours that increase the availability of suitable nests. Several studies reported ejection of host nestlings from nests by brood parasites; however, whether brood parasites do so to induce the host to re-nest and thus increase opportunities for future parasitism (i.e. \'farming\' behaviour) remains unclear. Here, we report observational evidence of farming behaviour by a common cuckoo Cuculus canorus female in a Daurian redstart Phoenicurus auroreus population: (1) the cuckoo destroyed a host nest by ejecting all nestlings, (2) the host then produced a new nest and (3) the cuckoo successfully parasitized the replacement nest. We suggest that farming behaviour may be more common, but often goes undetected because it requires intense nest monitoring.

Egg recognition and rejection are the most common and effective anti-parasitic strategies against avian brood parasitism in terms of maintaining stability over time and plasticity in response to environmental cues. Conversely, parasites have evolved multiple counter-adaptations to the anti-parasitic defenses of hosts. Among them, the crypsis hypothesis suggests that eggs that appear more cryptic in color and are closely matched to the environment are helping to counter the egg recognition strategy of the host. In this study, we investigated whether the egg recognition ability of Oriental reed warblers (Acrocephalus orientalis), a common host of common cuckoos (Cuculus canorus), changed during different reproductive stages by using model egg experiments. The effect of the crypsis hypothesis on the egg recognition ability of the hosts was also investigated by controlling the color contrast between the inside of the experimental nests and the model eggs. The results showed that the Oriental reed warbler retained strong egg recognition abilities, which were similar to the incubation stage (GLMMs: F 1,27 = 0.424, p = .521), even after entering the nestling stage and preferentially rejected model eggs with distinct contrasting colors (binomial test: Fisher\'s exact, p = .016). These results are consistent with the crypsis hypothesis. The present study suggests that the host retains a strong egg recognition ability even during the nestling stage and that cryptic-colored eggs that are closely matched with the breeding nest environment help counter the host\'s egg recognition abilities and increase the chances of successful parasitism by cuckoos. However, the effectiveness of cryptic egg may be weaker than mimic egg in countering egg recognition and rejection by hosts with open-cup nests.

Brood parasitism by cuckoos relies on manipulating hosts to raise their offspring and has evolved stunning adaptations to aid in their deception. The fact that cuckoos usually but not always, remove one or two host eggs while laying their eggs has been a longstanding focus of intensive research. However, the benefit of this behavior remains elusive. Moreover, the recently proposed help delivery hypothesis, predicting that egg removal by cuckoos may decrease the egg-laying duration in the parasitism process caused by biting action, lacks experimental verification. Therefore, in this study, we examined the effects of egg removal/biting on the egg-laying speed in the common cuckoo (Cuculus canorus) to experimentally test this hypothesis. We compared the duration of cuckoo egg-laying in empty nests, nests with host eggs, and nests with artificial blue stick models to test whether cuckoos biting an egg/stick can significantly hasten the egg-laying speed than no biting action. Our results showed that biting an egg or an object is associated with cuckoos laying approximately 37% faster than when they do not bite an egg or an object. This study provides the first experimental evidence for the help delivery hypothesis and demonstrates that when cuckoos bite eggs or other objects in the nest, they lay eggs more quickly and thereby avoid suffering the hosts\' injurious attack.

Comparative studies of egg recognition and rejection between various sympatric hosts provide insight into the coevolutionary history of the hosts and parasites, as well as the degree of antagonism between the species. Although buntings are widely considered to be a suitable host taxon for cuckoos, there has been relatively little research on this example of parasitism and host antiparasitic behaviour. Here we provided the first report on brood parasitism and egg recognition in three sympatric ground-nesting bunting hosts of the common cuckoo (Cuculus canorus), namely the yellow-throated bunting (Emberiza elegans), south rock bunting (E. yunnanensis), and crested bunting (E. lathami). The results show that for the five breeding seasons during 2018-2022, the parasitism rate by common cuckoos was 0.87% and 0.45% in yellow-throated buntings and south rock buntings, respectively, whereas the parasitism rate by an unidentified parasite was 4% during 2018-2023 in the crested bunting. The rejection rates of the three bunting hosts for blue non-mimetic eggs were 89.3%, 88.9%, and 100% for yellow-throated buntings, south rock buntings, and crested buntings, respectively. The rejection rates for red non-mimetic eggs by yellow-throated buntings and south rock buntings were lower at 76.9% and 82.4%, respectively. All three sympatric bunting hosts examined had high levels of egg recognition and egg rejection, suggesting that it may have been subjected to high parasitic history and that egg recognition ability was retained after the loss of parasitism, which needs to be further verified by future experiments.

It is a common occurrence for incubating female ground- or cavity-nesting birds to accidentally roll their own eggs out of the nest cup. To correctly roll eggs that are not in the nest cup back to the nest, birds must be able to discriminate between their eggs and things around their nests that could be easily confused for their eggs (e.g., egg-shaped stones or other birds\' eggs). Moreover, there may be a strong interaction between egg retrieval and egg rejection behaviors in the context of avian brood parasitism where recognition and rejection of alien eggs is an effective means for hosts to defend against cuckoo parasitism. It has been shown that green-backed tits (Parus monticolus) and Japanese tits (P. minor) in China have strong egg recognition ability and are able to reject nonmimetic eggs in the nest. Eggshell spots play an essential role in the egg rejection behavior of these two tit hosts. This study investigated the egg retrieval behavior of green-backed tits and Japanese tits by adding one experimental egg to the nest corner to explore whether eggshell spots also play a role in the process of egg retrieval. The results revealed significant differences in the retrieval rates of white-rumped munia (Lonchura striata) eggs painted with spots, Japanese tits\' own true eggs, and unpainted, pure white-white-rumped munia eggs. The retrieval rate of white-white-rumped munia eggs was significantly lower than that of spotted white-rumped munia eggs and Japanese tits\' own spotted eggs. For green-backed tits, the retrieval rate of spotted white-rumped munia eggs in the nest corner was significantly higher than that of pure white-white-rumped munia eggs, while the rejection rate of spotted white-rumped munia eggs was significantly lower than that of pure white-white-rumped munia eggs. These findings indicate that eggshell spots play a key role in the egg retrieval of green-backed tits and Japanese tits.

The arms race between brood parasites and their hosts provides a classic model to study coevolution. Hosts often reject the parasitic egg, and brood parasites should therefore select host nests in which the colour of the eggs best matches that of their own. Although this hypothesis has received some support, direct experimental evidence is still lacking. Here, we report on a study of Daurian redstarts, which show a distinct egg-colour dimorphism, with females laying either blue or pink eggs. Redstarts are often parasitized by common cuckoos, which lay light blue eggs. First, we showed that cuckoo eggs were more similar in spectral reflectance to the blue than to the pink redstart egg morph. Second, we report that the natural parasitism rate was higher in blue than in pink host clutches. Third, we performed a field experiment in which we presented a dummy clutch of each colour morph adjacent to active redstart nests. In this set-up, cuckoos almost always chose to parasitize a blue clutch. Our results demonstrate that cuckoos actively choose redstart nests in which the egg colour matches the colour of their own eggs. Our study thus provides direct experimental evidence in support of the egg matching hypothesis.

Urbanization is transforming ecosystems at a global scale and at an increasing rate,1,2 and its profound consequences for wildlife have been well documented.3,4,5,6 Understanding how animals thrive in the urban environment and how this environment affects (co-)evolutionary processes remains an important challenge.7 Urban environments can provide resources such as food or nest sites (e.g., cavities)10,8,9 and also reduce exposure to predators.11,12 For some species, urban environments may also affect susceptibility to brood parasitism,13,14 but this has never been tested experimentally. Here, we use a combination of field observations and experimental manipulations to show that Daurian redstarts, Phoenicurus auroreus, a common host of the common cuckoo, Cuculus canorus, nest in proximity to humans to avoid brood parasitism. First, redstarts were more likely to be parasitized with increasing distance to the nearest building. Second, redstarts adjusted their nesting location in response to a seasonally predictable change in the risk of brood parasitism. Third, experimentally simulating the presence of cuckoos during a period when they are naturally absent increased the likelihood that redstarts nested indoors or closer to human settlements. These findings suggest that redstarts actively choose to place their nest in the vicinity of a human residence as a defense against cuckoos. Our study exemplifies how animals take advantage of the urban environment by using it as a novel line of defense against detrimental interspecific interactions.

Nest defense is an effective strategy of hosts against parasites. Typically, hosts will aggressively attack brood parasites that approach or visit their nests, which can prevent the parasites from laying eggs or may even lead to the death of the parasites. Few previous studies have specifically reported such fatal cases involving brood parasites and have attributed the cause of death to either drowning or hypothermia after falling into the water following an attack from hosts. In this study, we recorded the process of multiple host individuals of the Oriental reed warbler (Acrocephalus orientalis) mobbing and attacking a female common cuckoo (Cuculus canorus) in the field. We discovered that the immediate cause of the cuckoo\'s death was the fatal physical damage resulting from the aggressive defense from the hosts, suggesting that frantic pecking and scratching by the hosts is the most proximate cause of mortality among egg-laying female cuckoos. This finding enhances our essential understanding of the effectiveness of host attacks.

Avian brood parasites leave parental care of their offspring to foster parents. Theory predicts that parasites should select for large host nests when they have sufficient available host nests at a given time. We developed an empirical experimental design to test cognitive ability of female cuckoos in nest size by studying nest choice of common cuckoos (Cuculus canorus) among nests of its Oriental reed warbler (Acrocephalus orientalis) hosts. We presented three groups of experimental nests: 1) nest dyads tied together including one large and one small artificial nest from reed leaves, 2) nest triads tied together used the old modified warbler\'s own nests including enlarged, reduced and medium-sized nests, and 3) nest dyads are similar to group 1, but not tied together to elicit parasitism by common cuckoos. We predict that cuckoos prefer larger nest than medium one, the next is smaller nest. Our findings showed that common cuckoo females generally prefer large nests over medium or small sized nests in all three experimental groups. Furthermore, cuckoo parasitism was significantly more common than in previous studies of the same warbler population, implying that larger, higher and more exposed host nests effectively increased the probability of cuckoo parasitism.