Abstract: We investigated the hypothesis that the common cuckoo Cuculus canorus selects host pairs of good phenotypic quality. As there is some evidence that cuckoos may select hosts within a population non-randomly based on external cues reflecting their foster abilities, we predicted that great reed warbler Acrocephalus arundinaceus pairs parasitized by the cuckoo would exhibit higher quality than unparasitized ones. To test this assumption, we evaluated two different parameters indicating host quality: body condition and characteristics of host eggs. We found that parasitized females showed significantly better body condition than unparasitized ones, and the model showed that the probability of being parasitized by the cuckoos increased with increasing body condition. Moreover, the likelihood of being parasitized by a cuckoo within the great reed warbler population increased with decreasing colour variability within clutches: parasitized females allocated costly blue pigments to eggshells more equally compared with unparasitized ones. Our study revealed that cuckoos parasitize great reed warbler females of higher quality, as reflected in host body condition and egg colour characteristics. In highly mimetic systems, cuckoos may choose to parasitize hosts with eggs displaying low intraclutch variation, both because this leads to reduced rejection and because these hosts are of high quality.

Abstract: Nest predation and brood parasitism are costly for nest owners, and natural selection should therefore favour the evolution of parental counterdefences. We addressed the question of whether Blackcaps Sylvia atricapilla change their incubation behaviour in response to various nest intruders and whether this adjustment matches the intensity of mobbing exhibited towards these intruders. Near focal nests, we successively exposed a dummy of a brood parasite, nest predator and an innocuous species. After the parents had responded, we removed the dummy and filmed their incubation. The most aggressive response towards the Common Cuckoo Cuculus canorus and high nest attendance after its disappearance indicated recognition of the brood parasite. Low-intensity response to the Jay Garrulus glandarius, together with reduced subsequent parental care, suggested that Blackcaps perceived it either as less deleterious at the egg stage than the Cuckoo or as a danger to themselves. Almost no aggression towards the Turtle Dove Streptopelia turtur, along with the resumption of incubation after its removal, implied that Blackcaps recognised it as harmless. In addition, we found that the level of aggression positively correlated with nest attendance, suggesting a link between the intensity of mobbing and subsequent parental care. Altogether, our results demonstrate that the issue of enemy recognition may be viewed as a complex of both aggressive and post-presentation behaviours.

Abstract: Nest protection against intruders is an indispensable component of avian parental care. In species with biparental care, both mates should evolve nest defence behaviour to increase their reproductive success. In most host–parasite systems, host females are predicted to have more important roles in nest defence against brood parasites, because they typically are primarily responsible for clutch incubation. Male antiparasitic behaviour, on the other hand, is often underestimated or even not considered at all. Here we investigated sex-specific roles in four aspects of great reed warbler (Acrocephalus arundinaceus) nest defence against a brood parasite—the cuckoo (Cuculus canorus), namely (1) mobbing, (2) nest attendance/guarding, (3) nest checking and (4) egg ejection. Using dummy experiments, simulating brood parasitism and by video-monitoring of host nests we found that males took the key roles in cuckoo mobbing and nest guarding, while females were responsible for nest checking and egg ejection behaviours. Such partitioning of parental roles may provide a comprehensive clutch protection against brood parasitism.

Abstract: After a population bottleneck, genetic drift can result in sudden and dramatic changes in allele frequencies that occur independently of selection. In such instances, many beneficial adaptations may be eliminated even if the population later increases again. Conversely, immigration may successfully counter the adverse effect of reduced population size, bringing new or lost alleles and replenishing thus genetic diversity of a reduced population. Extensive groundwater abstraction from the Azraq Oasis led to a considerable deterioration of the unique wetland ecosystem during the 1980s and most of the aquatic vegetation consequently died off in the early 1990s. At that time, numbers of local reed warblers (Acrocephalus scirpaceus) dramatically dropped to a few pairs. The numbers slightly increased again after wetland restoration efforts were begun. To infer possible consequences of the population collapse on genetic diversity, we genotyped 40 local breeders at 10 polymorphic microsatellite loci. After 15 years from the crash, the population showed no evidence for past genetic bottlenecks. The absence of reduced genetic diversity suggests that the population has largely benefited from gene flow. The study shows how immigration may sustain the viability of fragmented populations of migratory birds after restoration of their former breeding habitat.

Abstract: To investigate migratory connectivity in the Reed Warbler Acrocephalus scirpaceus, we analysed (1) all available sub-Saharan ringing recoveries and (2) stable isotopes in feathers grown in Africa sampled at 17 European breeding sites across a migratory divide. A cluster analysis of ringing recoveries showed remarkable connectivity between breeding and non-breeding grounds. Two main clusters represented populations taking the two main migratory routes [southwesterly (SW) and southeasterly (SE)]. Stable isotope analysis confirmed the separation of wintering areas of SW- and SE-migrating populations. Higher δ15N values in feathers of SE-migrating birds indicated that they occupied more xeric biome types. Values of δ13C that did not differ significantly among populations were higher than those from feathers of known European origin and indicated a C4 biome. Three populations with an unknown migratory direction were assigned to the SE-migrating populations on the basis of δ15N values.

Abstract: Three hundred and thirty-one Reed Warblers Acrocephalus scirpaceus (Hermann) were examined for chewing lice (Phthiraptera). Twenty-eight (8.5%) were parasitized with Menacanthus curuccae (Schrank, 1776). Mean intensity of infestation was 4.3 lice. Hence, the name Menacanthus eisenachensis Balát, 1981, currently applied to Menacanthus lice parasitizing Reed Warblers, is placed as a junior synonym of M. curuccae, with Reed Warbler as a new host for the latter species.

Abstract: Variation in appearance of parasitic and host eggs is an essential trait in the coevolutionary struggle between a brood parasite and its host. Here we examined the role of colour characteristics in the egg recognition in blackcaps, Sylvia atricapilla, assessed by (1) human vision and (2) spectrophotometry, including UV wavelengths. Since colour and spotting pattern within eggs are often highly variable, we also assessed three different parts of blackcap eggshell separately. We investigated the possibility that a lower intraclutch variation in the parasitized clutch would positively affect egg recognition. In addition, we predicted that egg rejection would depend on contrast between host and parasitic eggs. However, we found that neither intraclutch variation nor mimicry if assessed by humans had significant effect on host rejection. Multivariate analyses of spectrophotometric data revealed, however, that the likelihood of egg rejection significantly increased with decreasing intraclutch variation in blue chroma. Furthermore, probability of egg rejection significantly increased with decreasing contrast between parasitic and host eggs in UV wavelengths and with decreasing brightness of host eggs at the blunt egg pole. Accepted foreign eggs were significantly darker at the blunt pole than acceptors' own eggs. The paradox that rejected eggs had lower contrast could be explained by the fact that the contrast in UV spectra is associated with brightness of the blunt egg part. Our data suggest that colour characteristics of the blunt egg part may play a major role in the recognition of parasitic eggs in the blackcaps.

Abstract: Begging behavior of nestlings can signal both hunger and competitive ability. Studies of begging in evicting avian brood parasites exclude the influence of nestling competition and may provide new insights into the host–parasite conflict and the evolution of signaling. Apart from the begging call, common cuckoo Cuculus canorus nestlings use special vocal displays in the absence of their hosts, termed here host-absent vocalization (HAV). Since these conspicuous calls can increase the risk of predation and require energy, their costs should be balanced by some benefits, such as increased food provisioning. However, there has been no evidence that chicks convey information about their hunger by HAV. We therefore tested experimentally whether cuckoo chicks use HAV as an additional signal to enhance food-delivery rate by their hosts. We used playback of HAV recorded from cuckoo nestlings to determine whether their hosts, reed warblers Acrocephalus scirpaceus, increase their provisioning in response to an apparent increase in HAV. Older chicks spent more time in HAV than younger chicks, suggesting that HAV is not caused by inaccurate discrimination of host arrival stimuli. Negative correlation of HAV with feeding rate and mass gain between the two experiments suggested that hunger was the motivation of HAV. The playback experiment, however, did not prove that HAV affects host provisioning rate. We discuss possible reasons for this result and provide alternative explanations for HAV, such as creating a bond between the hosts and the parasitic young used later in the postfledging care.

Abstract: In some hosts of avian brood parasites, several populations apparently escape parasitism, while others are parasitized. Many migratory specialist brood parasites like common cuckoos, Cuculus canorus, experience a short breeding season, and in order to maintain local parasite populations host densities should be sufficiently high to allow efficient nest search. However, no studies have investigated the possible effect of host density on presence of cuckoo parasitism among populations of a single host species. Here, we investigated possible predictors of common cuckoo parasitism in 16 populations of reed warblers, Acrocephalus scirpaceus, across Europe. In more detail, we quantified the effect of host density, number of host breeding pairs, habitat type, mean distance to nearest cuckoo vantage point, predation rate and latitude on the presence of cuckoo parasitism while controlling for geographical distance among study populations. Host density was a powerful predictor of parasitism. We also found a less pronounced effect of habitat type on occurrence of parasitism, while the other variables did not explain why cuckoos utilize some reed warbler populations and not others. This is the first study focusing on patterns of common cuckoo-host interactions within a specific host species on a large geographic scale. The results indicate that if host density is below a specific threshold, cuckoo parasitism is absent regardless of the state of other potentially confounding variables.

Abstract: Much attention has been devoted to understanding the evolution of egg mimicry in avian brood parasites. The majority of studies have been based on human perception when scoring the mimicry of the parasitic egg. Surprisingly, there has been no detailed study on the recognition and sensitivity towards differently coloured parasitic eggs. We investigated effect of different colours of the experimental eggs measured by ultraviolet (UV)-visible reflectance spectrophotometry on rejection behaviour in the song thrush (Turdus philomelos). We carried out a set of experiments with four blue model eggs representing mimetic eggs, whereas six other colours represented nonmimetic eggs. Our results revealed that two colours originally designed as a mimetic were rejected at a high rate, whereas one group of the nonmimetic was accepted. A multiple regression model of absolute differences between song thrush and experimental eggs on rejection rate showed that the level of mimicry in the UV and green parts of the colour spectrum significantly influenced egg rejection in the song thrush. To our knowledge, this is the first detailed study showing that different colour perception by the birds can affect their responses towards the parasitic egg. These findings suggest that the combination of UV and visible ranges of the spectra plays a major role in the evolution of discrimination processes, as well as in the evolution of the mimicry of the parasitic egg.

Abstract: To evaluate host responses towards repeated brood parasitism we experimentally parasitized and continuously videotaped blackcap nests in two consecutive trials. The ejection of a foreign egg was the most common response (94.5%) in both trials, but desertion (4.1%) and acceptance (1.4%) also occurred. The general method of ejection was puncturing. In 9.8% of identified ejections, already punctured eggs stuck to the abdominal feathers of the incubating bird and were carried out of the nest. Females were responsible for the majority of ejections in both trials and their response time was significantly shorter than that of males. Blackcaps exhibited consistency in the sex responsible for egg ejection over the two trials; but in five (20.8%) experiments individuals changed their behaviour. Repeatability for host responses within the nest was very high. In ejections accomplished by the same bird, the response was significantly quicker in the second trial, indicating the presence of certain learning abilities. Our results suggest that cuckoo hosts are quite consistent in their responses towards parasitic eggs when parasitized repeatedly within one breeding attempt.

Abstract: We tested great reed warbler (Acrocephalus arundinaceus) discrimination against two common cuckoo (Cuculus canorus) female color morphs (gray and rufous) in two areas with different parasitism rates and proportions of the two morphs. Hosts recognized the two cuckoo morphs from a control, the feral pigeon (Columba livia), at Apaj, Hungary (where brood parasitism was heavy), whereas no significant differences among the models were recorded at Lužice, Czech Republic (where the parasitism rate was moderate). At Apaj, the hosts discriminated the rufous morph (which is slightly predominant there) better than the gray morph from the control. Between-site comparison (after controlling for background aggression) revealed that great reed warblers were more aggressive towards the rufous morph at Apaj than at Lužice, whereas their responses to the gray morph did not differ, corresponding with much higher between-site difference in the relative abundance of the rufous morph. Our results suggest that both local parasitism pressure and relative abundance of two female color morphs of a brood parasite may significantly influence host nest defenses.

Abstract: Blackcaps Sylvia atricapilla reject artificial cuckoo eggs, and their eggs vary little in appearance within clutches, whereas among clutches eggs vary considerably. Low variation within clutches facilitates discrimination of parasitic eggs, whereas high variation among clutches makes it harder for the cuckoo to mimic the eggs of a certain host species. These traits have most probably evolved as counteradaptations against brood parasitism by the common cuckoo Cuculus canorus, even though blackcaps are not regularly parasitised today. In this study, we investigated how fine-tuned the rejection of parasitic eggs is in this species by introducing three types of eggs into their nests: a real non-mimetic egg the approximate size of a cuckoo egg, an artificial mimetic egg the size of a cuckoo egg and a real conspecific egg. As the rejection frequency of both mimetic and non-mimetic artificial cuckoo eggs has been shown to be high in previous studies, the variation in rejection behaviour between individuals is low, indicating that most individuals within the population are able to reject parasitic eggs. Thus, we predict that (1) the intraclutch variation in egg appearance should be generally low in all individuals, and that (2) regarding conspecific eggs, rejection decisions should be highly dependent on the degree of mimicry between parasitic and host eggs. We found support for these predictions, which indicates that due to their highly sophisticated countermeasures against brood parasitism, blackcaps can probably be regarded as current winners of the arms race with the common cuckoo. Furthermore, the high and consistent rejection frequency of cuckoo eggs found throughout Europe for this species supports the spatial habitat structure hypothesis, which claims that woodland-nesting species breeding near trees, like blackcaps, presumably experienced a high level of parasitism throughout their range in the past and, therefore, their rejection behaviour, once evolved, spread rapidly to all populations.

Abstract:

Abstract: In a coevolutionary arms race between a brood parasite and its host, both species are expected to evolve adaptations and counteradaptations, such as egg mimicry and egg discrimination. The Common Whitethroat (Sylvia communis) is a regular Common Cuckoo (Cuculus canorus) host in some parts of its breeding range. We studied egg discrimination in the Whitethroat in the south-eastern part of the Czech Republic in an area where the Whitethroat is parasitised only occasionally. To investigate responses to parasitic eggs, in 34 nests either a non-mimetic or a mimetic (conspecific) egg was added. All 21 non-mimetic eggs were ejected. Of the 13 mimetic eggs, 7 were ejected and 6 eggs were accepted. No significant differences were found in intraclutch variation in egg appearance between acceptors and rejecters. Ejected mimetic eggs, however, showed greater contrast with host eggs than did the mimetic eggs that were accepted. Our experiments show that Whitethroats are persistent rejecters of alien eggs at the study site. This fact is discussed in the light of the host selection and host-parasite coevolution hypotheses.