Nest defence,enemy recognition and nest inspection behaviour of experimentally parasitized Reed Warblers Acrocephalus scirpaceus

Capsule Reed Warblers in a regularly parasitized population do not recognize Cuckoo Cuculus canorus as a special enemy and do not change their behaviour at nest immediately after being parasitized.

Aims To assess if an intruder near the nest influences the behaviour of the Cuckoo host.

Methods Host responses to Cuckoo, control Pigeon dummies and human intruder were observed. Host behaviour at 71 nests was video-recorded for 30 minutes at four experimental groups of nests: Cuckoo dummy, Cuckoo dummy + Cuckoo egg, Pigeon dummy, human intruder.

Results Reed Warblers did not respond differently to the Cuckoo and the control species. The experimental procedure had no significant effect on the behaviour of hosts during the study period. We were unable to find any differences in the time spent at the nest, clutch inspection behaviour and nest defence behaviour between morning and afternoon experimental groups. Our results do not support the hypothesis that afternoon laying by the Cuckoo is maintained by a selection pressure from the host. We observed no ejection or egg-pecking during the 30-min period after the experimental parasitism.

Conclusions Low aggression and non-specificity of host responses in our study area are in line with the fact that the Reed Warbler is an intermediate rejecter of Cuckoo eggs as expected from the spatial habitat structure hypothesis.     

Geographic variation in parasitism rates of two sympatric cuckoo hosts in China

Rates of brood parasitism vary extensively among host species and populations of a single host species. In this study, we documented and compared parasitism rates of two sympatric hosts, the Oriental Reed Warbler (Acrocephalus orientalis) and the Reed Parrotbill (Paradoxornis heudei), in three populations in China. We found that the Common Cuckoo (Cuculus canorus) is the only parasite using both the Oriental Reed Warbler and Reed Parrotbill as hosts, with a parasitism rate of 22.4%-34.3% and 0%-4.6%, respectively. The multiple parasitism rates were positively correlated with local parasitism rates across three geographic populations of Oriental Reed Warbler, which implies that higher pressure of parasitism lead to higher multiple parasitism rate. Furthermore, only one phenotype of cuckoo eggs was found in the nests of these two host species. Our results lead to two conclusions: (1) The Oriental Reed Warbler should be considered the major host of Common Cuckoo in our study sites; and (2) obligate parasitism on Oriental Reed Warbler by Common Cuckoo is specialized but flexible to some extent, i.e., using Reed Parrotbill as a secondary host. Further studies focusing on egg recognition and rejection behaviour of these two host species should be conducted to test our predictions.     

Common Cuckoos Cuculus canorus change their nest‐searching strategy according to the number of available host nests

In recent decades, numerous studies have examined factors affecting risk of host nest parasitism in well‐known avian host–parasite systems; however, little attention has been paid to the role of host nest availability. In accordance with other studies, we found that nest visibility, reed density and timing of breeding predicted brood parasitism of Great Reed Warblers Acrocephalus arundinaceus by the Common Cuckoo Cuculus canorus. More interestingly, hosts had a greater chance of escaping brood parasitism if nesting was synchronized. Cuckoo nest searching was governed primarily by nest visibility at high host‐nest density. However, even well‐concealed nests were likely to be parasitized during periods when just a few hosts were laying eggs, suggesting that Cuckoos adjust their nest‐searching strategy in relation to the availability of host nests. Our results demonstrate that host vulnerability to brood parasitism varies temporally and that Cuckoo females are able to optimize their nest‐searching strategy. Moreover, our study indicated that Cuckoos always manage to find at least some nests to parasitize. Thus, in this case, the co‐evolutionary arms race should take place mainly in the form of parasitic egg rejection rather than via frontline pre‐parasitism defence.     

Parasitierung und Brutverluste durch den Kuckuck (Cuculus canorus) bei Teich- und Sumpfrohrs?nger (Acrocephalus scirpaceus,A. palustris) in Mittel- und Westeuropa

Zusammenfassung Nach 52 brutbiologischen Untersuchungen und Angaben der Parasitierung durch den KuckuckCuculus canorus bei Teich- und SumpfrohrsängernAcrocephalus scirpaceus,A. palustris wird für Mittel- und Westeuropa eine durchschnittliche Parasitierung von 8,3±7,8 % bei Teich- und von 6,3±6,6 % bei Sumpfrohrsängern berechnet (Tab. 1). Wegen der hohen Streuung beträgt der Median beim Teichrohrsänger 9 % und beim Sumpfrohrsänger nur 1,2 %. Selbst unter der Annahme einer gewissen Unausgewohgenheit des Datenmateriales übertreffen diese Werte bei weitem die aller anderen häufigen Kuckuckswirte in Mittel- und Westeuropa. Die Parasitierung des Teichrohrsängers ist fast flächendeckend, beim Sumpfrohrsänger dagegen mehr lückig über die Region verteilt (Abb. 1). Die Parasitierungsfrequenz scheint im Osten und Süden geringer als im Nordwesten. Ihre hohe Brutbestandsdichte macht die beiden Rohrsänger zu idealen Kuckuckswirten. Es gibt Hinweise, daß die Parasitierung bei beiden Arten zunimmt (Abb. 2). Darüberhinaus werden auch zahlreiche Gelege von den Kuckucksweibchen geraubt. Diese Verluste können viermal so hoch sein wie die durch Parasitierung (s. auch Abb. 3). Beide Arten haben zahlreiche Abwehrmechanismen gegenüber dem Kuckuck, wobei die Eiablehnung durch den Sumpfrohrsänger viel heftiger und effektiver ist als beim Teichrohrsänger. Möglicherweise steht der Sumpfrohrsänger unter einem höheren selektiven Druck für erfolgreiches Brüten.

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Parasitism and egg losses due to the Cuckoo (Cuculus canorus) in Reed and Marsh Warblers (Acrocephalus scirpaceus,A. palustris) in central and western Europe

Summary Both Reed and Marsh Warblers are common Cuckoo hosts. In this paper the frequencies of Cuckoo parasitism in 52 studies dealing with breeding success as well as parasitism in both warblers over a wide range of central and western Europe are analysed. Average parasitism rates are 8.3±7.9 % in Reed and 6.3±6.6 % in Marsh Warblers (tab. 1). Taking the high variation coefficient into consideration the median of Reed Warbler parasitism is 9 % and of Marsh Warbler parasitism only 1.2 %. Due to the uneven distribution of the studies over the region analysed and due to the preference by the researchers for smaller study plots vs large habitats there might be some bias towards higher parasitism figures. Nevertheless, both warblers range on top of the list of Cuckoo hosts and have much higher parasitism rates than other common hosts. Parasitism of the Reed Warbler does occur over the whole region, whereas in the Marsh Warbler it is more unevenly and patchily distributed. Parasitism seems to be higher in the northwestern than in the eastern and southern parts of central and western Europe. Both warblers are ideal hosts due to their high population densities in good habitats. There are indications for an increase of parasitism in both species during the last 30–50 years. Besides parasitism, many clutches of both warblers are predated upon by female Cuckoos. The number of predated nests can be four times as high as the number parasitized (see also Fig. 3). Both species exhibit a broad array of counter-reactions against the Cuckoo. The egg rejection by Marsh Warblers is much stronger and much more effective than in the Reed Warbler. Probably Marsh Warblers are under a higher selection pressure for successful breeding than Reed Warblers or have a different strategy to avoid losses.

     

CuckooCuclus canorus parasitism onAcrocephalus Warblers in Southern Moravia in The Czech Republic

Summary The Reed Warbler,Acrocephalus scirpaceus, in the Lednice area, Southern Moravia in the Czech Republic, was parasitized by the Cuckoo,Cuculus canorus, at a rate of at least 18.0 %. The Cuckoo eggs showed poor mimesis with the Reed Warbler eggs, but showed a greater resemblance to the eggs of other species breeding in the area, including the Great Reed Warbler,A. arundinaceus. This latter species was also parasitized, but we did not find enough nests to obtain a reliable estimate for the rate of parasitism. The parasitized Reed Warblers rejected the Cuckoo eggs at a high rate (42.1 %) and therefore both the hatching success and the breeding success of the Cuckoo was considerably lower than shown by comparable results from Britain. On the background of these results (poor mimesis of the Cuckoo eggs and a high rejection rate by the hosts) the question of the degree of specialization versus generalism in the Cuckoo's host preference is discussed.

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Zusammenfassung Teichrohrsänger im Gebiet von Lednice (Südmähren, Tschechei) waren mindestens zu 18 % vom Kuckuck parasitiert. Die Kuckuckseier glichen jenen der Teichrohrsänger nur wenig, waren jedoch den Eiern anderer im Gebiet brütender Arten einschließlich des Drosselrohrsängers ähnlicher. Drosselrohrsänger waren auch parasitiert; wir fanden jedoch nicht genug Nester, um eine Parasitierungsrate ermitteln zu können. 42,1 % der Kuckuckseier wurden vom Teichrohrsänger nicht angenommen; daher waren Schlüpf- und Ausfliegerate des Kuckucks merklich geringer als in vergleichbaren Ergebnissen aus Großbritannien. Vor dem Hintergrund unserer Ergebnisse — geringe Angleichung der Kuckuckseier und hohe Ablehnungsrate durch die Wirtsart — wird die Frage von Spezialisierung und Generalismus in der Wirtswahl des Kuckucks diskutiert.

     

Experimental enlargement of nest size does not increase risk of predation or brood parasitism in the Great Reed Warbler Acrocephalus arundinaceus

We assessed whether nest size affects the probability of nest loss using dyads of large and small (large being twice the size of small) inactive Great Reed Warbler Acrocephalus arundinaceus nests placed at similar sites in Great Reed Warbler territories. Large nests were not predated significantly more frequently than small nests. Experimentally enlarged active Great Reed Warbler nests suffered non‐significantly higher predation compared with non‐manipulated control nests. Our experiments did not support the nest‐size hypothesis and suggested that nest size does not appear to be a factor affecting the risk of nest predation in this species. The probability of brood parasitism by the Common Cuckoo Cuculus canorus was also unaffected by experimental nest enlargement, supporting the commonly accepted hypothesis that the Common Cuckoo searches for suitable host nests by host activity during nest building rather than nest size.     

Breeding phenology,reproductive traits and apparent survival of sympatric Marsh Warbler Acrocephalus palustris and Blyth’s Reed Warbler Acrocephalus dumetorum

Capsule: Sympatric Marsh Warblers Acrocephalus palustris and Blyth’s Reed Warblers Acrocephalus dumetorum differ significantly in their life history traits.

Aims: To provide a direct comparison of demographic parameters among two sympatric populations of the closely related Marsh Warbler and Blyth’s Reed Warbler.

Methods: We examined breeding phenology and reproductive traits at a 25?ha study plot. We use program MARK to estimate daily nest survival and adult apparent survival rates.

Results: On average, Marsh Warblers laid the first eggs 3 days later than Blyth’s Reed Warblers. Mean clutch size in the Marsh Warbler was significantly lower than in the Blyth’s Reed Warbler. There are no significant differences between the two species for nest daily survival, duration of incubation and nestling periods. Apparent survival of adults was slightly higher in Marsh Warblers than in Blyth’s Reed Warblers.

Conclusion: Our findings suggest that two ecologically similar sympatric species differ significantly in terms of life history traits. We assume that observed differences could be the result of adaptations to environmental factors in the central parts of the species’ ranges or due to differences in mortality on migratory pathways or wintering grounds.     

Breeding ground correlates of the distribution and decline of the Common Cuckoo Cuculus canorus at two spatial scales

Many migratory bird species are undergoing population declines as a result of potentially multiple, interacting mechanisms. Understanding the environmental associations of spatial variation in population change can help tease out the likely mechanisms involved. Common Cuckoo Cuculus canorus populations have declined by 69% in England but increased by 33% in Scotland. The declines have mainly occurred in lowland agricultural landscapes, but their mechanisms are unknown. At both the local scale within the county of Devon (SE England) and at the national (UK) scale, we analysed the breeding season distribution of Cuckoos in relation to habitat variation, the abundance of host species and the abundance of moth species whose caterpillars are a key food of adult Cuckoos. At the local scale, we found that Cuckoos were more likely to be detected in areas with more semi‐natural habitat, more Meadow Pipits Anthus pratensis (but fewer Dunnocks Prunella modularis) and where, later in the summer, higher numbers of moths were captured whose larvae are Cuckoo prey. Nationally, Cuckoos have become more associated with upland heath characterized by the presence of Meadow Pipit hosts, and with wetland habitats occupied by Eurasian Reed Warbler Acrocephalus scirpaceus hosts. The core distribution of Cuckoos has shifted from south to north within the UK. By the end of 2009, the abundance of macro‐moth species identified as prey had also declined four times faster than that of species not known to be taken by Cuckoos. The abundance of these moths has shown the sharpest declines in grassland, arable and woodland habitats and has increased in semi‐natural habitats (heaths and rough grassland). Our study suggests that Cuckoos are likely to remain a very scarce bird in lowland agricultural landscapes without large‐scale changes in agricultural practices.     

The effect of Jacobin Cuckoo Clamator jacobinus parasitism on the body mass and survival of young in a new host species

The southern African subspecies of Jacobin Cuckoo Clamator jacobinus serratus is a brood parasite of a range of host species. While Jacobin Cuckoos do not evict host young, previous research has found that host young rarely survive the nestling period. Here we provide the first records of Jacobin Cuckoo parasitism of a new host species, the Southern Pied Babbler Turdoides bicolor. We investigate rates of brood parasitism and the survival of host young. The Southern Pied Babbler is one of the largest recorded hosts for Jacobin Cuckoos and, unusually, we find that host young tend to survive the nestling period and maintain similar body mass to host young in unparasitized broods. However, host young were less likely to survive to independence than young raised in unparasitized nests, suggesting a post‐fledging reproductive cost to hosts.     

Egg Rejection and Brain Size among Potential Hosts of the Common Cuckoo

Interspecific brood parasitism by the common cuckoo (Cuculus canorus) lowers host fitness, and has selected for discrimination and rejection of parasitic eggs in their commonly parasitized hosts. Cognitive demands needed to discriminate and reject cuckoo eggs may have led to augmentation of relative brain size among passerine hosts parasitized by cuckoos. This hypothesis predicts for across species positive relationships of brain size with rejection rate, host suitability and parasitism level. Here we test these predictions while controlling for phylogenetic, ecological and developmental factors known to affect brain size and egg rejection in a comparative study using the cuckoo and their hosts in Europe as a model system. Contrary to expected the rate of rejection of non‐mimetic cuckoo eggs covaried negatively with relative brain size across bird species. Either suitability as cuckoo host, which reflects long‐time duration of exposure to cuckoo parasitism, and level of parasitism, did not relate to brain size. Our results do not support the hypothesis that cuckoo parasitism was a main direct force affecting brain size variation across passerine hosts.     

Song in two cryptic species: comparative analysis of Large-billed Reed Warblers Acrocephalus orinus and Blyth’s Reed Warblers Acrocephalus dumetorum

Capsule: Songs of Large-billed Reed Warblers Acrocephalus orinus and Blyth’s Reed Warblers Acrocephalus dumetorum differed in quantitative parameters. Blyth’s Reed Warbler used different modes of singing on breeding grounds and migration stopovers.

Aims: To compare the songs of two cryptic species of reed warblers. To compare Blyth’s Reed Warbler songs in different parts of the breeding range and on migration stopovers in central Asia. To investigate the status of Blyth’s Reed Warblers in central Asia.

Methods: We analysed song recordings of individual singing males and conducted field observations of singing behaviour at migration stopover and breeding sites. The status of Blyth’s Reed Warblers in central Asia was reviewed from the historical Russian literature and other sources.

Results: We found differences between six out of eight variables in songs of Large-billed and Blyth’s Reed Warblers on breeding sites. Blyth’s Reed Warbler songs recorded on migration stopovers showed differences in five variables from Large-billed Reed Warblers and in six variables from breeding Blyth’s Reed Warblers. On migration stopovers, Blyth’s Reed Warblers sang actively but did not stay more than 1–2 days. The evidence suggests that Blyth’s Reed Warbler does not breed in central Asia.

Conclusion: Songs of Large-billed and Blyth’s Reed Warblers could be distinguished by quantitative analysis. Blyth’s Reed Warblers used different modes of singing on breeding sites and migration stopovers. We believe vocalizations of Blyth’s Reed Warblers on migration to be plastic song.     

When should Common Cuckoos Cuculus canorus lay their eggs in host nests?

Capsule Brood parasitic Common Cuckoo Cuculus canorus chicks hatch earlier than the nestlings of their Great Reed Warbler Acrocephalus arundinaceus hosts, but hatching priority is less consistent when Cuckoo eggs are laid after the onset of host incubation.

Aim To reveal by field observations what the optimal stage is for Cuckoos to lay their eggs in relation to the host laying cycle to ensure prior hatching of the parasitic chicks.

Methods We monitored the hatching of Cuckoo chicks in relation to the hosts’ laying stage at which the Cuckoo eggs appeared and also monitored host incubation behaviour.

Results Great Reed Warblers incubated more on day 5 after the host's onset of laying relative to day 3. All Cuckoo eggs hatched earlier than hosts when they were laid prior to the onset of host incubation (day 4). Cuckoo eggs also maintained hatching priority in about 2/3 of the nests when laid on days 5–6.

Conclusions Most Cuckoo eggs are laid prior to the onset of host incubation and this, together with other adaptive mechanisms, ensures the prior hatching of Cuckoo eggs. Cuckoo eggs laid after the onset of incubation lose the advantage of prior hatching in approximately 30% of nests.     

Common Cuckoo Cuculus canorus and host behaviour at Reed Warbler Acrocephalus scirpaceus nests

This study is based on continuous video recordings made at 53 Reed Warbler Acrocephalus scirpaceus nests each day during the laying period. Egg-laying by the Common Cuckoo Cuculus canorus was recorded in 14 (26.4%) of these nests. By analysing the activity of the host birds around and at the nest, we found that this is probably not the only cue used by the Common Cuckoo when locating suitable nests to parasitize. Furthermore, in most cases there was no significant difference between the length of time the host birds spent at the nest in the morning and afternoon, thus providing little support for the hypothesis that the Common Cuckoo lays in the afternoon because it is less likely to be seen by the nest owners then. Parasitized Reed Warblers rejected the Common Cuckoo egg more frequently when they observed the parasite at their nests. However, contrary to what should be expected, most Common Cuckoos laid their eggs in the presence of the host(s), and in general their egg-laying behaviour (for example duration of stay at the nest) was less secretive than described earlier. When partially depredating host clutches, Cuckoos showed the same behavioural pattern at parasitized and unparasitized nests, indicating that the latter may act as a potential reserve for egg-laying.     

The spatial habitat structure of host populations explains the pattern of rejection behavior in hosts and parasitic adaptations in cuckoos

In this article we present tentative support for predictionsderived from a spatial habitat structure hypothesis arguingthat common cuckoos Cuculus canorus, the most common obligatebrood parasite in Europe, only breed in areas where they haveaccess to vantage points in trees. Thus, species in which somepopulations breed near trees while other populations breed farther from trees have a different cuckoo—host population dynamicthan species in which all populations always breed in the vicinityof trees. Parasitism rate, mimicry of brood parasite eggs withthose of the hosts, and rejection behavior of hosts varieswith the host breeding habitat. Cuckoos are best adapted toexploit species in which some populations breed near trees while other populations breed in open areas because such hosts arenot always accessible to cuckoos, and thus gene flow amongunparasitized and parasitized populations delays the evolutionof host adaptations. Adaptive behavior in cuckoos as well asin their hosts can be predicted from the spatial habitat structurehypothesis.     

How to hatch from an egg of great structural strength. A study of the Common Cuckoo

Brood parasitism represents a unique mode of avian reproduction that requires a number of adaptations. For example, to reduce chances of puncture ejection of their eggs by small hosts, brood parasites may have been selected for laying eggs of unusually great structural strength. However, great structural strength of eggshells should hinder hatching. The goals of our study were to establish if chicks of the Common Cuckoo Cuculus canorus have more difficulty with hatching out of their strong eggs than chicks of species with eggs of similar size, and whether they possess any mechanisms facilitating hatching. To achieve these goals, we compared hatching pattern and selected body characteristics of chicks of the Common Cuckoo with those of another altricial species with eggs of a similar size, the Great Reed Warbler Acrocephalus arundinaceus . Although the rate of pecking was similar in the two species, the Common Cuckoo chicks started pecking earlier in relation to their emergence and consequently required more time and a greater cumulative number of pecks for breaking open their eggs than did young Great Reed Warblers. The two species also differed with respect to the pattern of opening their shells; in contrast to the warbler chicks, which enlarged the original pip circularly, the cuckoo chicks opened the egg by systematically creating a long narrow slit until they emerged. Finally, our study of hatched young revealed several differences; the Cuckoo hatchlings were significantly heavier, had a longer forearm, and their egg tooth was located significantly farther from the tip of the beak. The edge used for cutting through the shell was also significantly longer than that of hatchling Great Reed Warblers. To conclude, our data suggest that hatching is more difficult for a Cuckoo than for a Great Reed Warbler and that Cuckoos possess several mechanisms to overcome the problems of hatching from a structurally strong egg.     

外寄生性花绒寄甲的寄生选择及其发育表现

"选择-表现"假说认为,成虫应该选择有利于子代发育的高品质寄主,但在寄主选择中,除了寄主品质外,其他因素也可能影响寄主选择决策。寄主选择研究通常以成虫为对象,而对那些初龄幼虫选择寄主的寄生性昆虫很少关注。以1龄幼虫积极搜寻寄主的寄生性花绒寄甲为模式生物,采用双选试验设计,观察了花绒寄甲初孵幼虫在不同体重青杨天牛幼虫之间、在已被寄生与健康的黄粉虫蛹之间的寄生选择性;然后采用回归设计,观察了花绒寄甲寄生若干不同体重的青杨天牛幼虫后的发育表现。研究结果表明,花绒寄甲1龄幼虫对体型较大的青杨天牛幼虫的选择偏好显著大于对体型较小的寄主幼虫的选择,选择大体型幼虫的比值比是选择小体型幼虫的4.55倍;对已被寄生的寄主黄粉虫蛹的选择偏好显著大于对健康寄主蛹的选择,选择已被寄生寄主的比值比是选择健康寄主的12.57倍。寄生青杨天牛幼虫的花绒寄甲幼虫发育历期平均为11.49 d、蛹历期为26.67 d、幼虫发育至成虫的羽化率50%,这些发育表现与寄生时青杨天牛幼虫的体重没有显著关系。但刚羽化寄甲成虫体重与寄生时寄主的体重存在显著的正直线关系:寄生时的寄主体重每增大0.01 g,羽化出的寄甲成虫体重增大近0.08%;方差分析寄甲成虫体重在不同寄主体重水平之间的差异表明,从体型较大寄主中羽化的寄甲成虫体重显著大于从体型较小寄主中羽化的成虫。研究结果说明,花绒寄甲初孵幼虫在寄主选择决策时,在寄主体型大小与被寄生状态之间可能采取折衷对策,而且对体型大小不同的寄主选择与子代发育适合度表现存在一致性,从而支持"选择-表现"假说。     

To eject or to abandon? Life history traits of hosts and parasites interact to influence the fitness payoffs of alternative anti‐parasite strategies

Hosts either tolerate avian brood parasitism or reject it by ejecting parasitic eggs, as seen in most rejecter hosts of common cuckoos, Cuculus canorus, or by abandoning parasitized clutches, as seen in most rejecter hosts of brown‐headed cowbirds, Molothrus ater. What explains consistent variation between alternative rejection behaviours of hosts within the same species and across species when exposed to different types of parasites? Life history theory predicts that when parasites decrease the fitness of host offspring, but not the future reproductive success of host adults, optimal clutch size should decrease. Consistent with this prediction, evolutionarily old cowbird hosts, but not cuckoo hosts, have lower clutch sizes than related rarely‐ or newly parasitized species. We constructed a mathematical model to calculate the fitness payoffs of egg ejector vs. nest abandoner hosts to determine if various aspects of host life history traits and brood parasites’ virulence on adult and young host fitness differentially influence the payoffs of alternative host defences. These calculations showed that in general egg ejection was a superior anti‐parasite strategy to nest abandonment. Yet, increasing parasitism rates and increasing fitness values of hosts’ eggs in both currently parasitized and future replacement nests led to switch points in fitness payoffs in favour of nest abandonment. Nonetheless, nest abandonment became selectively more favourable only at lower clutch sizes and only when hosts faced parasitism by a cowbird‐ rather than a cuckoo‐type brood parasite. We suggest that, in addition to evolutionary lag and gape‐size limitation, our estimated fitness differences based on life history trait variation provide new insights for the consistent differences observed in the anti‐parasite rejection strategies between many cuckoo‐ and cowbird‐hosts.     

Survival of Reed Warbler Acrocephalus scirpaceus clutches in relation to nest position

Nest survival was studied in relation to nest position of 164 nests of the Reed Warbler Acrocephalus scirpaceus in the Southeastern part of the Czech Republic in 1993. Breeding was successful in 91 (55.5%) cases, whereas 40 (24.4%) were predated, 19 (11.6%) were parasitized by the Cuckoo Cuculus canorus and 14 (8.5%) were destroyed by other causes. Of the predated nests, small mammals accounted for 26 (65.0%) and unknown predators for 14 (35.0%) nests. Rates of predation and parasitism varied in relation to a series of habitat factors and Reed Warblers avoided nest sites most vulnerable to predation or parasitism. Concealed places in the vegetation far away from trees were the safest nest sites.     

Post-fledging interactions between the Common Cuckoo Cuculus canorus and its cavity-nesting Common Redstart Phoenicurus phoenicurus host

Brood parasite–host interactions during the incubation and nestling stages have been well studied, but the post-fledging period remains virtually unknown. Using radiotracking, we provide the first detailed data on post-fledging interactions between the Common Cuckoo Cuculus canorus and its only regular cavity-nesting host, the Common Redstart Phoenicurus phoenicurus. Cuckoos raised alone (‘solitary’) fledged at higher mass, with higher wing and tarsus length and started to fly at a younger age than Cuckoos raised alongside young Redstarts (‘mixed’). However, a further 23 fledging and post-fledging parameters measured at five pre-determined times (fledging, first-flight, predation, starvation, independence) did not differ between solitary and mixed Cuckoos. In addition, none of the parameters measured during the post-fledging period (growth, dispersal distances, number of flights) differed between solitary and mixed Cuckoos. Redstart fledglings from non-parasitized broods (‘solitary’) showed generally similar fledging and post-fledging parameters to fledglings reared alongside a Cuckoo (‘mixed’). Surprisingly, there were no significant differences in post-fledging predation rate, starvation or overall survival rates between mixed and solitary Cuckoos or mixed and solitary Redstarts. Thus, during the post-fledging period, mixed Cuckoo fledglings successfully compensated for the poorer performance experienced during the nestling stage whereas mixed and solitary Redstarts did not differ in any measured parameters. This suggests that the regular occurrence of mixed broods in this host–parasite system – which is unique among the many Cuckoo hosts – is evolutionarily stable for both hosts and parasites.     

Responses of potential hosts of Asian cuckoos to experimental parasitism

In the arms race between avian brood parasites and their hosts, several adaptations and counter‐adaptations have evolved. The most prominent host defence is rejection of parasitic eggs. We experimentally parasitized nests of 10 potential host species breeding in sympatry with four different cuckoo species in an area in Bangladesh using differently coloured model eggs to test host responses. In four species we introduced both mimetic and non‐mimetic eggs. Black Drongos Dicrurus macrocercus, hosts of the Indian Cuckoo Cuculus micropterus, rejected all model eggs. Common Mynas Acridotheres tristis and Jungle Babblers Turdoides striata accepted all eggs regardless of mimicry. These two species are parasitized by Asian Koels Eudynamys scolopaceus, Common Hawk‐cuckoo Hierococcyx varius and, in the case of Jungle Babblers, Jacobin Cuckoos Clamator jacobinus. Pied Mynas Gracupica contra, with no records of parasitism in our study area, also accepted all eggs regardless of mimicry. In the six remaining species, all of which lay spotted eggs, we introduced only non‐mimetic eggs. Black‐hooded Orioles Oriolus xanthornus rejected all model eggs, even though we have found no records of natural parasitism. Long‐tailed Shrikes Lanius schach and House Crows Corvus splendens, hosts of Asian Koels, rejected 75 and 9.1% of model eggs, respectively. Large‐billed Crows Corvus macrorhynchos, apparently not used as hosts in our study area, accepted all blue but rejected all brown model eggs. Oriental Magpie‐Robins Copsychus saularis and Red‐vented Bulbuls Pycnonotus cafer accepted all non‐mimetic model eggs. In Black Drongos, Long‐tailed Shrikes and Black‐hooded Orioles, all model eggs were ejected within 24 h of introduction. The results show considerable variation in egg rejection rates among various species, providing baseline data for further investigation of co‐evolutionary interactions between brood parasites and hosts in this region.