Decoupled Acoustic and Visual Components in the Multimodal Signals of the Common Cuckoo (Cuculus canorus)

Because of its parasitic habits, reproduction costs of the common cuckoo (Cuculus canorus) are mostly spent in pre-laying activities. Female costs are limited to searching host nests and laying eggs, whereas, males spend time in performing intense vocal displays, possibly with territorial purpose. This last aspect, together with a sexual plumage dimorphism, points to both intra- and inter-sexual selections operating within this species. One element triggering sexual selection is a differential fitness accrued by different phenotypes. Before analyzing possible sexual selection mechanisms operating in cuckoos, it is therefore necessary to verify whether there is a variability among male secondary characters by describing and quantifying them. Here we aimed to quantitatively characterize the main two potential candidates of sexual selection traits, i.e., calls and displays, shown by males at perches. During the 2019 breeding season, in a site within the Po Plain, we both audio and video recorded cuckoo males at five different perches. We analyzed acoustic variables as well as display sequences searching for potential correlations. We found a significant variation among calls that could be clustered into four vocal types. We also found that no visual displays were associated with vocal displays; cuckoo males were either vocal and motionless or soundless and active. We discuss our results under the perspective of the potential value of sexual selection in brood parasites and its role in its parasitic habit.

Host eyes, brains and their brood parasites

1. Brood parasites interact with their hosts for exploitation of host parental abilities and the associated resources. This results in coevolutionary interactions of hosts and parasites. 2. A prime example of such a common specialist brood parasite is the common cuckoo Cuculus canorus and its host races. Hosts use their cognitive abilities to identify parasites and vice versa for their ability to discriminate among potential hosts. 3. We predicted that parasites with relatively large brains for their body size should be more successful at avoiding their hosts, and that hosts with small brains for their body size should more often be exploited by parasites. We also predicted that hosts with relatively large eyes for their body size would have superior discriminatory abilities allowing for superior discrimination against brood parasites. Finally, we predicted that visual ability of specific cuckoo hosts would have evolved exaggerated visual ability as estimated from the relative size of their optic tectum would have resulted in such hosts being more successful as reflected by their higher rate of parasitism. 4. Interspecific variation in size of brain, eye, optic tectum, telencephalon and cerebellum were consistent with these predictions.

Testing the Interspecific Function of Female Common Cuckoo “Bubbling” Call

Female common cuckoo (Cuculus canorous) predator-like “bubbling” calls distract host parental attention and reduce the egg rejection rate. Such “bubbling” calls are also frequently used to attract males and deter territorial rivals in intraspecies contact, and these calls are an ancestral character in many cuckoo species. Although hosts have had sufficient time to become familiar with this call and evolve anti-parasitic strategies, why are the hosts fooled by this “bubbling” call? We propose two hypotheses. The first hypothesis proposes that call variation reduces the opportunity for host species to correctly assess cuckoo tricks. In contrast, the second hypothesis proposes that the cost of behavior may prevent the antiparasitic strategy from evolving. In the study, we tested the prerequisites of these hypotheses, by investigating whether cuckoo calls vary during the day and testing whether the predator-like calls suppress bird activities. Based on field recordings from three different areas, we found high overlap in the calls generated during different periods. Oriental great reed warblers (Acrocephalus orientalis), a host species, did not show different responses toward the playback of female common cuckoo calls generated before noon or afternoon. Based on bird count data, we found that predator-like call playback is insufficient for suppressing bird activities. Therefore, none of the prerequisites were supported by our field data. We discuss the potential reasons for our findings and hope to inspire more research examining female cuckoo vocalizations.

Behavioral and Acoustic Responses of the Oriental Reed Warbler (Acrocephalus orientalis), at Egg and Nestling Stages, to the Common Cuckoo (Cuculus canorus)

Cuckoo nest parasites lay eggs in host nests and thereby transfer all reproduction costs to the hosts. This greatly reduces host fitness. Parasitism has selected for the evolution of anti-parasitic strategies in hosts, including nest defense. The dynamic risk assessment hypothesis holds that nest parasitism only threatens the nests during the egg stage, so hosts should reduce the level of defense against nest parasites after the egg stage. We studied the behavioral and acoustic responses of oriental reed warblers (Acrocephalus orientalis), during both the egg and nestling stages, toward the common cuckoo (Cuculus canorus), sparrowhawks (Accipiter nisus) and oriental turtle doves (Streptopelia orientalis). A. orientalis can visually distinguish cuckoos from sparrowhawks and doves, indicating that hawk mimicry did not work for the cuckoos. The behavioral response of hosts in the nestling stage was stronger than in the egg stage, which supports the offspring value hypothesis and suggests that cuckoos may also act as nest predators. However, there was no difference in the alarm calls A. orientalis produce in response to different invaders, indicating that different types of alarm calls may not contain specific information.

Call rate in Common Cuckoos does not predict body size and responses to conspecific playbacks

The brood parasitic Common Cuckoo Cuculus canorus is best known for its two-note “cu-coo” call which is almost continuously uttered by male during the breeding season and can be heard across long distances in the field. Although the informative value of the cuckoo call was intensively investigated recently, it is still not clear whether call characteristic(s) indicate any of the phenotypic traits of the respective vocalising individuals. To fill this gap, we studied whether the call rate of male cuckoos (i.e., the number of calls uttered per unit of time) provides information on their body size, which might be a relevant trait during intrasexual territorial conflicts. We captured free-living male cuckoos and measured their body size parameters (mass, wing, tail and tarsus lengths). Each subject was then radio-tagged, released, and its individual “cu-coo” calls were recorded soon after that in the field. The results showed that none of the body size parameters covaried statistically with the call rates of individual male Common Cuckoos. In addition, we experimentally tested whether the “cu-coo” call rates affect behavioural responses of cuckoos using playbacks of either a quicker or a slower paced call than the calls with natural rates. Cuckoos responded similarly to both types of experimental playback treatments by approaching the speaker with statistically similar levels of responses as when presented with calls at the natural rate. We conclude that male Common Cuckoos do not advertise reliable information acoustically regarding their body size, and so, cuckoo calls are neither useful to characterize cuckoos’ phenotypic traits directly nor to indicate environmental quality indirectly.

Personality of hosts and their brood parasites

Brood parasites such as the common cuckoo Cuculus canorus exploit the parental abilities of their hosts, Hosts avoid brood parasitism and predation by showing specific behaviour such as loss of feathers, emission of fear screams and contact calls, displaying wriggle behavior to avoid hosts or potential prey, pecking at hosts and prey and expressing tonic immobility (showing behaviour like feigning death or rapid escape from predators and brood parasites). These aspects of escape behavior are consistent for individuals but also among sites, seasons and years. Escape behaviour expressed in response to a broad range of cuckoo hosts and prey are consistently used against capture by humans, but also hosts and brood parasites and predators and their prey. An interspecific comparative phylogenetic analysis of escape behavior by hosts and their brood parasites and prey and their predators revealed evidence of consistent behaviour when encountering potential parasites or predators. We hypothesize that personality axes such as those ranging from fearfulness to being bold, and from neophobic to curiosity response in brood parasites constitute important components of defense against brood parasitism that reduces the overall risk of parasitism.

Does experimentally simulated presence of a common cuckoo (Cuculus canorus) affect egg rejection and breeding success in the red-backed shrike (Lanius collurio)?

Providing artificial eggs is a commonly used technique to understand brood parasitism, mainly by the common cuckoo (Cuculus canorus). However, the presence of a cuckoo egg in the host nest would also require an earlier physical presence of the common cuckoo within the host territory. During our study of the red-backed shrike (Lanius collurio), we tested two experimental approaches: (1) providing an artificial “cuckoo” egg in shrike nests and (2) additionally placing a stuffed common cuckoo with a male call close to the shrike nest. We expected that the shrikes subject to the additional common cuckoo call stimuli would be more sensitive to brood parasitism and demonstrate a higher egg rejection rate. In the years 2017–2018, in two locations in Poland, a total of 130 red-backed shrike nests were divided into two categories: in 66 we added only an artificial egg, and in the remaining 64 we added not only the egg, but also presented a stuffed, calling common cuckoo. Shrikes reacted more strongly if the stuffed common cuckoo was present. However, only 13 incidences of egg acceptance were noted, with no significant differences between the locations, experimental treatments or their interaction. Analysis of breeding success revealed significant differences between the locations, between experimental treatments and their interaction, which suggests a strong location effect. The red-backed shrike is an efficient rejector of foreign eggs. It would be interesting to see how similar tests affect hosts that have much higher rates of brood parasitism and egg acceptance.