Two main mechanisms of egg rejection, the main defence of hosts against brood parasites, have been described: ejection and desertion. Desertion of the parasitized nest is much more costly and is usually exhibited by small-sized host species unable to remove the parasitic egg. However, nest desertion is frequently assumed to be an anti-parasite strategy even in medium or large-sized host species. Th is assumption should be considered with caution because: 1) large-sized hosts able to eject the parasitic egg should eject it rather than desert the nest, and 2) breeding birds may desert their nests in response to diff erent disturbances other than brood parasitism. Th is problem is especially important in the common blackbird Turdus merula because this is a medium-sized species, potential host of the common cuckoo Cuculus canorus , in which desertion has been frequently reported as a response to cuckoo egg models. Here, we seek to determine whether nest desertion can be considered a response unequivocally directed to the parasitic egg in medium-sized hosts using the blackbird as the study species. In an experimental study in which we have manipulated levels of mimicry and size of experimental eggs, we have found that both colour (mimetic and non-mimetic; at least for human vision) and size (small, medium, and large) signifi cantly aff ected ejection rates but not nest desertion rates. In fact, although large eggs disproportionally provoked nest desertion more frequently than did small or medium-sized eggs, cuckoo-sized parasitic eggs were not deserted allowing us to conclude that desertion is unlikely to be an adaptive response to brood parasitism at least for this species.

Brood parasitism frequently leads to a total loss of host fitness, which selects for the evolution of defensive traits in host species. Experimental studies have demonstrated that recognition and rejection of the parasite egg is the most common and efficient defence used by host species. Egg-recognition experiments have advanced our knowledge of the evolutionary and coevolutionary implications of egg recognition and rejection. However, our understanding of the proximate mechanisms underlying both processes remains poor. Egg rejection is a complex behavioural process consisting of three stages: egg recognition, the decision whether or not to reject the putative parasitic egg and the act of ejection itself. We have used the blackbird (Turdus merula) as a model species to explore the relationship between egg recognition and the act of egg ejection. We have manipulated the two main characteristics of parasitic eggs affecting egg ejection in this grasp-ejector species: the degree of colour mimicry (mimetic and non-mimetic, which mainly affects the egg-recognition stage of the egg-rejection process) and egg size (small, medium and large, which affects the decision to eject), while maintaining a control group of non-parasitized nests. The behaviour of the female when confronted with an experimental egg was filmed using a video camera. Our results show that egg touching is an indication of egg recognition and demonstrate that blackbirds recognized (i.e., touched) non-mimetic experimental eggs significantly more than mimetic eggs. However, twenty per cent of the experimental eggs were touched but not subsequently ejected, which confirms that egg recognition does not necessarily mean egg ejection and that accepting parasitic eggs, at least sometimes, is the consequence of acceptance decisions. Regarding proximate mechanisms, our results show that the delay in egg ejection is not only due to recognition problems as usually suggested, given that experimental eggs are not touched significantly more often. Thus, the delay in egg ejection is mainly the consequence of a delay in the decision to eject, probably triggered by mechanical constraints imposed by eggs that are harder to eject (i.e. larger). Our results offer important information on the relationships between recognition and ejection and contribute to a better understanding of host defences against brood parasites.

Many host species have evolved sophisticated defences to mitigate the high fitness costs imposed by brood parasitism. Even though the physiological mechanisms behind such defences can offer important insights into the evolutionary relationship between brood parasites and hosts, they have received little attention so far. Hormones play a critical role in the regulation of bird reproduction, which make them a key element when investigating the physiological effects of brood parasitism on hosts. Here, we experimentally parasitized Eurasian blackbird (Turdus merula) nests with non-mimetic eggs to study its impact on the hormonal levels (corticosterone and prolactin) of females during incubation, as well as the magnitude of the response to the standardized stress protocol in parasitized and non-parasitized individuals. Parasitized females had higher baseline corticosterone levels and showed a poorer body condition than non-parasitized birds, while we found no differences for prolactin levels. Both parasitized and non-parasitized females responded to the standardized-stress protocol with a significant increase in corticosterone levels. However, the decrease in prolactin after the standardized stress protocol was significantly more pronounced in parasitized individuals. Our results suggest that the presence of a non-mimetic parasitic egg involves a stressful situation for hosts, negatively affecting the physical state of parasitized females. Unaffected prolactin levels of parasitized individuals could explain the absence of nest desertion found in this species in response to parasitism. Finally, both hormones were not correlated in blackbirds, confirming that their combined study provides valuable pieces of information on the endocrine mechanisms underlying behavioural responses in animals, including hosts of brood parasites. Significance statementPhysiological mechanisms behind avian brood parasitism remain unclear. In this study, we assessed the effect of experimental parasitism on the hormonal profiles of hosts. We found that the presence of a non-mimetic egg in the nest modified baseline corticosterone levels, but not prolactin levels, of parasitized females and negatively impacted their body condition. Moreover, experimental parasitism affected the prolactin response to stress. These results expand previous information on the endocrine consequences of brood parasitism at other stages of the breeding cycle (nestling and fledgling stage) and might shed light on the hormonal mechanisms that underlie the host response against parasitic eggs. Communicated by M. Leonard

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.

The post-fledging period is a critical phase for juvenile survival, and parental care provided during this period is a key component of avian reproductive performance. Very little is known about the relationships between foster parents and fledglings of brood parasites. Here, we present the results of a 5-year study about the relationships between fledglings of the non-evictor brood parasitic great spotted cuckoo (Clamator glandarius) and its magpie (Pica pica) foster parents. Sometimes, great spotted cuckoo and magpie nestlings from the same nest can fledge successfully, but most often parasitic nestlings outcompete host nestlings and only cuckoos leave the nest. We have studied several aspects of cuckoo post-fledging performance (i.e. feeding behaviour, parental defence and fledgling survival) in experimental nests in which only cuckoos or both magpie and cuckoo nestlings survived until leaving the nest. The results indicate that great spotted cuckoo fledglings reared in mixed broods together with magpie nestlings were disadvantaged by magpie adults with respect to feeding patterns. Fledgling cuckoos reared in mixed broods were fed less frequently than those reared in only cuckoo broods, and magpie adults approached less frequently to feed cuckoos from mixed broods than cuckoos from only cuckoo broods. These results imply that the presence of host's own nestlings for comparison may be a crucial clue favouring the evolution of fledgling discrimination; and furthermore, that the risk of discrimination at the fledgling stage probably is an important selection pressure driving the evolution of the arms race between brood parasites and their hosts