Egg Recognition Ability Research Articles

Breeding pairs with color aberrations in oriental reed warblers.

In 2017, one pair of Oriental reed warblers (Acrocephalus orientalis) with color aberrations was found in Yongnianwa National Wetland Park, Hebei, China. The female bird exhibited white feathers on the head, neck, and upper back, and the base of the beak was flesh-red in color. The male had a few feathers on the outer edges of the left and right primary wing coverts that were white, which was determined to be leucism after analysis. The breeding pairs laid their first egg on May 29, with a clutch size of four eggs. After an incubation period of 13 days, two chicks hatched on June 13, 2017. The nest was found empty on June 20 when the chicks were 7 days old and before fledging age; therefore, it was presumed that the chicks had been predated. A white parrot egg was added to the nest during the incubation period to test the egg recognition ability of breeding pairs and was successfully rejected. To the best of our knowledge, this is the first report of color aberrations in the Oriental reed warbler, and we found that this color aberrations did not affect some reproductive and antiparasitic behaviors of the birds, but whether it affects their breeding success needs to be further studied.

Oriental reed warblers retain strong egg recognition abilities during the nestling stage.

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 and egg recognition in three bunting hosts of the cuckoos.

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.

Eggshell spots are an important cue for the egg retrieval behavior in two tit species.

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.

Egg rejection and egg recognition mechanism in a Chinese Azure-winged Magpie (Cyanopica cyanus) population

Recognition and rejection of foreign eggs are effective defense of hosts against brood parasitism. However, brood parasitism can impose various selection pressures on different geographic populations of the same host species. In a multiple cuckoo system in China, Azure-winged Magpies (Cyanopica cyanus) are parasitized by both Indian Cuckoos (Cuculus micropterus) and Asian Koels (Eudynamys scolopaceus). In this study, egg recognition ability and recognition mechanism of the Azure-winged Magpie were investigated using a population in Fusong, southeastern Jilin, China. The results showed that 55.6% (20/36) of the Azure-winged Magpies correctly rejected quail (Coturnix japonica) eggs in their nests, while 13.9% (5/36) of the individuals experienced rejection costs by wrongly rejecting their own eggs. Azure-winged Magpies could accurately reject the experimental eggs when the number of such eggs in the nests was the same as that of the magpie eggs. However, Azure-winged Magpies do not recognize and reject conspecific eggs (0/28). The present study indicates that the Azure-winged Magpie has moderate egg recognition ability toward non-mimetic quail eggs and shows a true recognition mechanism with rejecting foreign eggs by accurately recognizing their own eggs. However, they cannot recognize conspecific eggs.

Egg recognition and nestling discrimination in the Crested Myna (Acridotheres cristatellus): Size matters

Most studies exploring abilities of hosts to detect brood parasitism are based on detecting colour and/or pattern differences among parasitic and host eggs or nestlings, while only few were focused on size differences. True recognition and recognition by discordancy are used to explain cognitive mechanisms of host egg recognition; however, only a few studies have found that hosts use recognition by discordancy. This study investigated: 1) whether egg and nestling recognitions in the Crested Myna (Acridotheres cristatellus) are based on size cues; 2) whether the egg cognitive mechanism is recognition by discordancy based on size cues; and 3) whether the longer the experiment time, the higher the egg recognition rate. Our results showed that the Crested Myna uses egg or nestling size as a recognition cue while the egg and nestling colour and patterning are not associated with egg or nestling rejection, thus the cognitive mechanism of egg recognition in the Crested Myna is recognition by discordancy based on egg size cues. Furthermore, there is a rejection delay in time of egg rejection behaviour of the Crested Myna. Therefore, we suggest that the periodicity of egg rejection experiments could be appropriately extended, especially for species with relatively low egg recognition ability.

Egg rejection and egg recognition mechanisms in Oriental Reed Warblers

BackgroundNest parasitism by cuckoos (Cuculus spp.) results in enormous reproductive failure and forces hosts to evolve antiparasitic strategies, i.e., recognition of own eggs and rejection of cuckoo eggs. There are often sexual conflicts between male and female individuals in the expression of antiparasitic behavior due to the differences in reproductive inputs and division of labor.MethodsBy adding a foreign egg made of blue soft clay to the host nest during early incubation period in the field, and by removing several host eggs and adding experimental eggs to control the proportion of two egg types in the nest, we examined egg rejection ability, egg recognition mechanism and sexual difference in egg rejection of the Oriental Reed Warbler (Acrocephalus orientalis), one of the major hosts of Common Cuckoos (Cuculus canorus).ResultsOur results indicated that Oriental Reed Warblers can recognize and reject nearly 100% (73/75) of the non-mimetic eggs made of blue soft clay, and they could reject foreign eggs with 100% accuracy, regardless of the ratio of experimental eggs and its own eggs in the nest. Furthermore, all cases of egg rejections recorded by videos were only carried out by females.ConclusionsOriental Reed Warblers have a high egg recognition ability and show a true recognition mechanism. Only female warblers perform egg rejection, suggesting that the sex for host egg incubation seems to play an important role in the evolution of egg recognition mechanisms.

Cryptic eggs are rejected less frequently by a cuckoo host.

Obligate brood parasitism is associated with huge reproduction costs, forcing hosts to evolve various anti-parasitic strategies against brood parasites, among which egg recognition and rejection is the most effective defense strategy. According to the crypsis hypothesis, non-mimetic yet cryptic eggs in a nest can also deceive their hosts and eventually be accepted. To validate this hypothesis, we conducted field experiments on Oriental reed warblers (Acrocephalus orientalis), a common host for common cuckoos (Cuculus canorus). We firstly tested the egg recognition and rejection abilities of Oriental reed warblers, using black and white model eggs in natural nests. Then we designed a comparison test where the cryptic effects of the two groups of experimental eggs were different. We manipulated the nest lining color and added relatively cryptic and bright model eggs to test warblers' rejection behaviors against cryptic and bright foreign eggs. The results showed that warblers have strong egg recognition and rejection abilities. There is a significant tendency for warblers to prefer to peck and reject relatively distinguishable foreign eggs, which supports the crypsis hypothesis. These findings indicate that even in the host-parasite system of open nests, parasitic eggs that are cryptic enough are prevented from being discovered and rejected by the host, and thus obtain the possibility of successful parasitism.

Best of a bad job or masters of illusion: Do nest light conditions make the eggs of brood parasitic brown‐headed cowbirds (Molothrus ater) more similar to the eggs of their hosts?

AbstractBrown‐headed cowbirds (Molothrus ater) are generalist obligate brood parasites, laying in the nest of nearly 300 avian species, and successfully parasitizing well over 100 host species. Cowbird eggs are generally considered non‐mimetic, although some have suggested that cowbird eggs resemble several of their host species’ eggs. To date, no investigation has examined the level of avian‐perceived similarity between cowbird and diverse host eggs in the contexts of light characteristics at the nest and the visual system of the relevant viewer. Because the cowbird exploits a wide range of species that lay in a variety of nest types, hosts view these eggs under an array of light conditions which could facilitate or hinder egg discrimination. When considering the visual system of the relevant viewers and the light conditions at their nest, we found that the coloration of cowbird eggs was more similar to host than non‐host species’ eggs. Host responses (whether they accept or reject cowbird eggs) were not statistically different when hosts perceived a large chromatic difference between their own eggs and the cowbird's eggs. Instead, we found that host responses were predicted by the degree to which nesting light conditions facilitated color similarity between host and cowbird eggs, such that hosts typically nesting under light conditions where this color discrimination task was more challenging were more likely to reject cowbird eggs. This suggests that the nesting light environment may have selected for increased coevolved egg recognition abilities in a suite of cowbird host species, even in the absence of parasitic egg color mimicry.

Potential cuckoo hosts have similar egg rejection rates to parasitized host species

BackgroundThrush species are rarely parasitized by cuckoos, but many have a strong egg recognition ability. To date, there is a limited understanding of the relationship between host egg rejection and cuckoo parasitism rate.MethodsBy using egg experiments in the field, we compared egg rejection between two non-parasitized potential host species and two parasitized hosts of cuckoos in the same region.ResultsThe White-bellied Redstart (Luscinia phoenicuroides), a host of the Common Cuckoo (Cuculus canorus), rejected 66.6% of blue model eggs; the Elliot’s Laughingthrush (Trochalopteron elliotii), a host of the Large Hawk Cuckoo (Hierococcyx sparverioides), rejected 25% of blue model eggs and 46.1% of white model eggs; and the Chestnut Thrush (Turdus rubrocanus) and the Chinese Thrush (T. mupinensis), in which cuckoo parasitism has not been recorded, rejected 41.1 and 83.3% of blue model eggs, respectively. There were no significant differences in the egg rejection among them, although the Chinese Thrush showed the highest rate of egg rejection.ConclusionsThis study indicates that the egg recognition ability of cuckoo hosts has no correlation with the actual parasitism rate of cuckoos. We suggest that the egg recognition ability of the two potential host species may have been retained from a parasitic history with the cuckoo, while the two common host species have developed their egg rejection abilities due to current parasitism pressure. In addition, our study highlights the importance of the multi-cuckoo parasite system for better understanding the selection pressure of parasitism on the evolution of host egg recognition abilities.

Egg recognition and brain size in a cuckoo host

The evolution of animal brain size and cognitive ability is a topic of central significance in evolutionary ecology. Interspecific brood parasitism imposes severe selection pressures on hosts favoring the evolution of cuckoo egg recognition and rejection. However, recognizing and rejecting foreign parasitic eggs are enormous cognitive challenges for cuckoo hosts, which might select for an increase in brain size in birds with this capacity. To explore the association between cuckoo parasitism and the evolution of brain size in cinereous tits (Parus cinereus), we used two types of experimental parasitic eggs, real mimetic white-rumped munia (Lonchura striata) eggs and non-mimetic blue model eggs, to test the egg recognition ability of female cinereous tits, thereby comparing brain size variation among individuals that were able to recognize foreign eggs and those that lacked this ability. Interestingly, our results however did not support the prediction that cuckoo parasitism selects for an increase in brain size of host birds, since brain size of egg rejecters was not significantly larger than that of accepters. Hence, this study suggested that the evolution of cognitive ability did not allow recognition of foreign eggs by female cinereous tits. That was the case despite the evolution of a larger brain may have allowed for a reduction in the cost of brood parasitism by cuckoos.

Egg rejection and egg recognition mechanism of chestnut thrushes (Turdus rubrocanus)

Recognizing and rejecting foreign eggs is one of the most effective anti-parasite strategies for hosts in avian brood parasitism. Studies have shown that most cuckoo (Cuculus spp.) hosts have evolved egg recognition abilities. Although some open-nesting birds, especially thrushes belonging to the family Turdidae, are rarely parasitized by cuckoos, they still have high egg recognition ability. This evolutionary selection pressure on egg recognition is currently controversial. Previous studies on egg recognition of thrushes have mainly been carried out in Europe in a single-cuckoo system. In this study, chestnut thrushes (Turdus rubrocanus), which are distributed in a multiple-cuckoo system in China, were used to test their egg rejection and egg recognition mechanism. Our results showed that chestnut thrushes had a rejection rate of 54 % for non-mimetic blue model eggs, showing moderate egg recognition ability. Their egg recognition was true recognition, which relied on a memory template; chestnut thrushes could accurately reject foreign eggs in their nests. This study added the second case to report the egg recognition mechanism of thrushes in the Turdidae family and showed that the evolution of egg recognition ability of chestnut thrushes was likely a retained anti-parasitic strategy because of being parasitized by cuckoos in the past.

Model eggs fail to detect egg recognition in host populations after brood parasitism is relaxed

BackgroundObligate 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.ResultsHere, 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.ConclusionsOur 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.

Egg recognition in Cinereous Tits (Parus cinereus): eggshell spots matter

BackgroundBrood parasitic birds such as cuckoos (Cuculus spp.) can reduce their host’s reproductive success. Such selection pressure on the hosts has driven the evolution of defense behaviors such as egg rejection against cuckoo parasitism. Studies have shown that Cinereous Tits (Parus cinereus) in China have a good ability for recognizing foreign eggs. However, it is unclear whether egg spots play a role in egg recognition. The aims of our study were to investigate the egg recognition ability of two Cinereous Tit populations in China and to explore the role of spots in egg recognition.MethodsTo test the effect of eggshell spots on egg recognition, pure white eggs of the White-rumped Munia (Lonchura striata) and eggs of White-rumped Munia painted with red brown spots were used to simulate experimental parasitism.ResultsEgg experiments showed that Cinereous Tits rejected 51.5% of pure white eggs of the White-rumped Munia, but only 14.3% of spotted eggs of the White-rumped Munia. There was a significant difference in egg recognition and rejection rate between the two egg types.ConclusionsWe conclude that eggshell spots on Cinereous Tit eggs had a signaling function and may be essential to tits for recognizing and rejecting parasitic eggs.

Egg recognition and chick discrimination in colonial breeding birds

Theory predicts that parents will not raise unrelated offsprings. For colonial breeding birds, evolving an ability to recognize their own eggs and chicks can prevent misdirecting parental behaviour. To verify this hypothesis, egg recognition experiments were performed on colonial breeding Chinese pond herons (Ardeola bacchus) and cattle egrets (Bubulcus ibis) in Chengmai and Lingao, tropical Hainan Island, China. Furthermore, a chick discrimination experiment was conducted on Chinese pond herons. In our study area, we did not record conspecific or interspecific brood parasitism in the two heron species. The results showed that Chinese pond herons had similar recognition rates for the four types of experimental eggs and lacked recognition ability. Cattle egrets had variable abilities to recognize different types of foreign eggs and have low egg recognition ability. Chinese pond herons cannot discriminate foreign chicks. Therefore, we suggest that the two species of colonial breeding herons may lack or have low egg recognition ability, and Chinese pond herons show no nestling recognition ability.

Egg-recognition abilities in non-incubating males: implications for the evolution of anti-parasitic host defenses

In the field of brood parasitism, it has been traditionally assumed that only the incubating sex rejects parasitic eggs, but this assumption has been rarely explored despite its important implications for the evolutionary relationship between brood parasites and hosts. Here, we used information on previous egg-rejection experiments to explore the recognition abilities of both males and females of Eurasian blackbirds Turdus merula towards experimental eggs with a variable degree of mimicry. We found that both sexes recognized non-mimetic eggs, supporting the idea that visits to the nest can favor the evolution of rejection abilities. In contrast, only females recognized mimetic eggs, indicating that although recognition abilities can evolve in both sexes, they are subsequently refined in females probably due to their more frequent interaction with parasitic eggs. Clutch size affected nest attendance since females, but not males, spent more time at the nest and visited it more frequently in larger clutches. Finally, our recordings showed that blackbird males are able not only to recognize, but also to eject parasitic eggs. Our results provide new insights into the main anti-parasitic defense in birds, egg rejection, and highlight the need of considering the role of the non-incubating sex in egg-rejection studies. Given the high costs associated to avian brood parasitism, both sexes are expected to evolve anti-parasitic defenses. However, in those species in which only females incubate, females have traditionally been assumed to be the responsible for egg rejection. Here, using the Eurasian blackbird (Turdus merula), we investigated the existence of egg-rejection abilities in non-incubating males and compared them to those exhibited by females. We found that males can recognize non-mimetic eggs, although their recognition abilities were less fine-tuned compared to females, who also recognized mimetic eggs. Even though females were the responsible for most documented egg-ejection events, recordings confirmed that males could also be involved in egg ejection, which could have important implication for the evolution of anti-parasitic defenses in host populations.

Russet Sparrows spot alien chicks from their nests

BackgroundIn coevolutionary interactions between brood parasites and their hosts, host parents are under strong selection to evolve defenses against parasitism. Egg rejection is an efficient and common defense against parasitism, although some apparently suitable hosts do not reject cuckoo eggs.MethodsSparrows Ploceidae are widespread throughout the Old World, and they have a suitable diet for rearing cuckoos, but still they are rarely exploited by brood parasites. To solve such puzzle, we conducted artificial parasitism and cross-fostering experiments in Russet Sparrow (Passer cinnamomeus).ResultsThe present study showed that Russet Sparrows have no egg recognition ability, but recognize their own nestlings and eject alien chicks or starve them to death. They may use visual cues in chick recognition, although they accept sister species Tree Sparrow (Passer montanus).ConclusionsBy rejecting nestlings of foreign species, Russet Sparrows have succeeded to escape from the brood parasitism by cuckoos and other parasites. Our studies shed light on the puzzle why some species are not utilized by cuckoo parasites as hosts.

Egg rejection in blackbirds Turdus merula: a by-product of conspecific parasitism or successful resistance against interspecific brood parasites?

BackgroundTraditional theory assumes that egg recognition and rejection abilities arise as a response against interspecific brood parasitism (IBP). However, rejection also appears in some species that are currently not exploited by interspecific parasites, such as Turdus thrushes. Recent evidences suggest that rejection abilities evolved in these species as a response to conspecific brood parasitism (CBP). To test these two alternative hypotheses, we performed an experimental study by parasitizing nests of the common blackbird (Turdus merula) with conspecifics or heterospecific eggs under different risk of parasitism (presence of interspecific or conspecific parasites near the nest). Common blackbird is a potential host of the common cuckoo (Cuculus canorus) but suffers low levels of CBP too.ResultsWe found that blackbirds were able to recognize and eject heterospecific eggs at high rates whereas most of conspecifics eggs were not recognized and, therefore, accepted. Ejection rates of conspecific eggs did not exceed 13 %, even in situations of high risk of CBP (blackbird female placed near the nest), which contradict the main prediction derived from the CBP hypothesis. Conversely, ejection rates of experimental eggs simulating IBP were much higher (80–100 %). Furthermore, female blackbirds were more aggressive towards cuckoos than towards blackbird dummies.ConclusionsOur results considered together support the IBP hypothesis, indicating that recognition and rejection of parasitic eggs in blackbirds have probably evolved due to previous cuckoo parasitism. The current absence of IBP in blackbirds may be due to the highly efficient rejection abilities in this species. Thus, these abilities have been retained in absence of brood parasitism as a consequence of the low costs involved for blackbirds, resulting in a successful resistance against interspecific brood parasitism.

Contrasting egg recognition between European and Asian populations of tree sparrows (Passer montanus)

Although many biological phenotypes are generally regarded as consistent across the distributional range of a species, some traits such as egg discrimination behavior have been shown to display extensive intraspecific variation as a response to selection from brood parasitism. We investigated the egg recognition ability in an Asian population of tree sparrows (Passer montanus), and we compared that with the ability to recognize and reject intraspecific foreign eggs in a population in Europe. Extensive artificial parasitism with model eggs and real eggs of eight sympatric birds that vary in background color and markings revealed that egg recognition capacity is completely absent in this Asian population of tree sparrows. This result contrasts with previous studies in European populations showing extensive ability for discriminating between own and foreign eggs. Different evolutionary equilibria or differences in the risk of conspecific parasitism may account for differences in egg discrimination ability between European and Asian populations of tree sparrows.

Nest sanitation behavior in hirundines as a pre-adaptation to egg rejection to counter brood parasitism.

Previous studies suggested that nest sanitation behavior may have been a pre-adaptation from which egg rejection of brood parasite eggs evolved. We tested this hypothesis in two swallow species, the red-rumped swallow (Cecropis daurica) and the barn swallow (Hirundo rustica). Our results indicated that the red-rumped swallow, which is an accepter of foreign eggs, rejected a low percentage of non-egg-shaped objects and did so less often than the barn swallow, which is an intermediate rejecter of foreign eggs. Furthermore, the egg rejection rates of the barn swallow increased with the increase in rejection rates of non-egg-shaped objects among different populations. These results showed that nest cleaning behavior could have evolved into a means of reducing the costs of brood parasitism, suggesting that egg recognition ability has evolved from recognition of non-egg-shaped objects. This finding advances our understanding of the evolution of egg recognition behavior in birds.