Provisioning of nestling Cuckoos Cuculus canorus by Reed Warbler Acrocephalus scirpaceus hosts

Reed Warblers Acrocephalus scirpaceus fostering a single nestling Cuckoo Cuculus canorus bring food to it at roughly the same rate as they do to an average-sized brood (three or four) of their own young. The food brought, mostly flies and beetles, is also similar. We conclude that the Cuckoo does not provide a supernormal stimulus. We show that Reed Warblers, given experimentally-enlarged broods of seven or eight, can substantially increase their feeding rate. This raises the question of why the young Cuckoo does not exploit this 'spare' feeding capacity of the Reed Warbler hosts. We offer three explanations', (i) that the increased begging necessary would attract predators, (ii) that the young Cuckoo is unable to grow faster, and (iii) that it would not be to the advantage of a young Cuckoo, dependent on its foster parents for about 5 weeks (cf. 3 weeks for Reed Warbler young), to provoke a feeding rate that the warblers could not sustain.     

European Cuckoo Cuculus canorus parasitism and host's rejection behaviour in a heavily parasitized Great Reed Warbler Acrocephalus arundinaceus population

An unusually high frequency (64%) of European Cuckoo Cuculus canorus parasitism was found in Great Reed Warbler Acrocephalus arundinaceus clutches in central Hungary. Sixty-four per cent of the parasitized clutches contained one Cuckoo egg, 23% contained two, 10% had three and 3% had four. This means that 58% of the Cuckoo eggs were found in multiply parasitized clutches. In multiple parasitism the laying second Cuckoo removed an egg from the clutch randomly, so preferred neither the host eggs, nor the concurrent Cuckoo egg. Host response towards the parasitic eggs showed 66% acceptance, 12% ejection, 20% desertion and 2% egg burial. We found great variation in both the host and the parasitic egg colour and pattern. This reduces the chance that the parasitic egg's appearance matched that of the hosts' but, in spite of this, almost perfect mimesis was found in 28% of the Cuckoo eggs. Poorly mimetic Cuckoo eggs were more frequently rejected by Great Reed Warblers than parasite eggs that were very similar to the host eggs. This high level of mimicry sometimes makes it difficult for the observer to identify the parasitic egg, especially when it is similar in size to the host eggs. It is also difficult for the host, as shown by the relatively high recognition error and ejection cost.     

Errors in egg-laying by female Common Cuckoo Cuculus canorus in nests of its common host

Dozens of studies have documented that brood parasites are well adapted to a brood parasitic lifestyle but not all parasitism events are successful. Co-evolution between brood parasites and their hosts is a dynamic process so it is reasonable to expect that a female brood parasite may commit errors during egg deposition by laying her eggs outside the laying period of the host, with consequent impacts on her fitness. Using an extensive dataset from a long-term study, we evaluated egg-laying patterns and errors related to the timing of egg-laying in the Common Cuckoo Cuculus canorus (hereafter ‘Cuckoo’). Specifically, we tested whether the Cuckoo avoids laying before or on the day of host clutch initiation to reduce the risk of rejection of parasitic eggs, whether laying errors will be more frequent in periods with a lack of active host nests, and whether the laying errors will be more frequent in periods with intense Cuckoo parasitism and a consequent lack of suitable host nests. We found that about one-third of Cuckoo eggs were laid on the host clutch initiation day or 1 day before, and the percentage of Cuckoo eggs laid decreased thereafter. Surprisingly, the probability of Cuckoo egg acceptance by the hosts was not affected by the egg-laying stage of the host clutch. Errors in the timing of egg-laying with fatal consequences (i.e. those precluding Cuckoo hatching because of laying in incubated or deserted clutches) were recorded in about 5% of cases. Only laying date of a Cuckoo egg had a significant effect on the probability of errors, which increased during the breeding season. This may be related to the higher number of deserted and incubated host nests at the site at the end of the breeding season. Errors in egg-laying may be attributed to young and inexperienced females but also impaired body condition or intraspecific competition may cause this behaviour. Future studies, which will test these possible explanations, will help to understand better the mechanism of co-evolutionary arms races and differences between host specialist and generalist brood parasites in various host–parasite systems.     

Responses of breeding Reed Warblers Acrocephalus scirpaceus to mounts of Sparrowhawk Accipiter nisus, Cuckoo Cuculus canorus and Jay Garrulus glandarius

Mounts of a Cuckoo Cuculus canorus , a Sparrowhawk Accipiter nisus and a Jay Garrulus glandarius were presented at nests of Reed Warblers Acrocephalus scirpaceus at all breeding stages. Response strength could best be classified according to approach distance (to the mount) and vocalizations used. Warblers reacted much more aggressively to a mount on the nest than to one 3 m adjacent; they approached much more closely, were less likely to sing but often gave a frenzied-sounding rasp (which was never given to an adjacent mount). Reaction was more aggressive to a Cuckoo, which was often attacked, than to a Sparrowhawk, of which they were wary; response to a Jay was intermediate. Reaction was more aggressive after the clutch had been completed than before. After the brood had fledged, parents still responded strongly to the Jay and Sparrowhawk, but responded to the Cuckoo only briefly before ignoring it. These results are consistent with the Reproductive Value-Stimulus Value hypothesis. They also demonstrate that Reed Warblers can tell Cuckoos from Sparrowhawks, as can many non-host species. The close resemblance of these two species is often suggested to be mimicry of the Sparrowhawk by the Cuckoo but, if this is so, it is not successful. Alternatively, the similarity could be mimicry to reduce predation on the Cuckoos themselves, a chance resemblance, or due to similar selective forces acting on both species, as each would presumably benefit from reduced conspicuousness.     

The common cuckoo Cuculus canorus is not locally adapted to its reed warbler Acrocephalus scirpaceus host

The obligate avian brood parasitic common cuckoo Cuculus canorus comprises different strains of females that specialize on particular host species by laying eggs of a constant type that often mimics those of the host. Whether cuckoos are locally adapted for mimicking populations of the hosts on which they are specialized has never been investigated. In this study, we first explored the possibility of local adaptation in cuckoo egg mimicry over a geographical mosaic of selection exerted by one of its main European hosts, the reed warbler Acrocephalus scirpaceus. Secondly, we investigated whether cuckoos inhabiting reed warbler populations with a broad number of alternative suitable hosts at hand were less locally adapted. Cuckoo eggs showed different degrees of mimicry to different reed warbler populations. However, cuckoo eggs did not match the egg phenotypes of their local host population better than eggs of other host populations, indicating that cuckoos were not locally adapted for mimicry on reed warblers. Interestingly, cuckoos exploiting reed warblers in populations with a relatively larger number of co-occurring cuckoo gentes showed lower than average levels of local adaptation in egg volume. Our results suggest that cuckoo local adaptation might be prevented when different cuckoo populations exploit more or fewer different host species, with gene flow or frequent host switches breaking down local adaptation where many host races co-occur.     

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.     

大杜鹃雌鸟鸣声的日节律

本文描述了三个地区大杜鹃(Cuculus canorus)雌鸟鸣声的日节律。通过在北京野鸭湖湿地公园(2017年5月7日至7月8日)、辽宁辽河口自然保护区(2018年6月28日至7月29日)和吉林大岗子林场(2018年5月17日至7月10日)分别布设自动录音机10台、10台和8台,连续记录繁殖季大杜鹃的鸣声。共录到2 777次大杜鹃雌鸟的鸣叫。其中,1 431次来自北京野鸭湖湿地公园,1 222次来自辽宁辽河口自然保护区,124次来自吉林大岗子林场。野鸭湖湿地公园和辽河口自然保护区的大杜鹃雌鸟在日出时段出现鸣叫的高峰,而大岗子林场的大杜鹃在日出和日落时段各有一个鸣叫的高峰。通过比较大杜鹃雌鸟鸣声的日节律和大杜鹃产卵时间的日节律,本研究认为在上述地区大杜鹃雌鸟鸣叫的功能更可能是宣示领域和吸引异性,而非恐吓宿主。本研究有助于进一步了解大杜鹃雌鸟鸣声的功能。     

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.     

The Large-billed Reed Warbler Acrocephalus orinus revisited

The Large-billed Reed Warbler Acrocephalus orinus is known only from the type specimen, collected in Himachal Pradesh, India, in 1867. The specimen is poorly prepared, and it has been suggested that it could represent an isolated form of the Clamorous Reed Warbler A. stentoreus or an aberrant Blyth's Reed Warbler A. dumetorum . We tested the affinity of A. orinus by (1) re-examining the morphology of the type specimen and (2) amplifying and sequencing a portion of its mitochondrial cytochrome b gene. Both the morphological and the mitochondrial analyses showed the specimen to be similar to dumetorum , but distinct enough to qualify as a species of its own. Relative to dumetorum , it has a more rounded wing, longer bill, longer and more graduated tail with more pointed tail feathers, and larger claws. The divergence in mitochondrial DNA between orinus and dumetorum was 7.8%, well above the value expected between subspecies. A. orinus is smaller than any of the forms of A. stentoreus or the related Australian Reed Warbler A. australis . It has a somewhat longer first primary, more pointed tail feathers and paler, less robust feet and claws. DNA comparison places it in the clade of small unstreaked Acrocephalus warblers, and apart from the clade of large unstreaked warblers that contains stentoreus and australis .     

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.     

Effects of mate removal on the behaviour and reproductive success of Reed Warblers Acrocephalus scirpaceus

The Reed Warbler Acrocephalus scirpaceus is a largely monogamous insectivorous passerine in which males and females provide equal care by day to eggs and chicks. Polygyny occurs occasionally, with males leaving one female unaided. When females were temporarily removed from three recently-completed clutches their males deserted and resumed the high song levels typical of unmated males. Males may desert either because they are physically incapable of incubation or because the energy expenditure needed for a male to return to an equivalent stage in the breeding cycle is much lower than for a female to do so. Widowed females ( n = 7 ), however, continued the breeding attempt alone, with similar incubation levels but higher provisioning rates than those of control females. In three out of four mid-season broods raised by lone females all fertile eggs were reared to healthy fledglings (in the fourth brood the female died), while only one of four late-season nests produced any fledglings (which were underweight). Late-season control nests were as successful as earlier ones. Loss of male help led to starvation of chicks, but caused no adverse effects during incubation. This explains the small changes in widows' sitting levels during incubation, but much greater effects after hatching. Females may need male help to rear late broods (but not early broods) as days are shorter and food is scarcer. Males may normally help at nests, even those in the mid-season, because in stressful spells (even for a few days) such help is vital for successful breeding but in good periods it costs the male little.     

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.     

Rejection of artificial cuckoo (Cuculus canorus) eggs in relation to variation in egg appearance among reed warblers (Acrocephalus scirpaceus)

Passerines that are exposed to brood parasitism can evolve reduced intraclutch variation in egg appearance to facilitate recognition and rejection of the parasitic egg. This has been shown to be true for European passerine species that are assumed to have participated in an evolutionary arms race with the cuckoo (Cuculus canorus). However, few investigations have been carried out with the aim of finding out whether there is a relationship between these two traits within a species. In this study, we compare the level of intraclutch variation in egg appearance and the rejection of an unlike parasitic egg within a population of reed warblers (Acrocephalus scirpaceus) in the south-eastern part of the Czech Republic. We parasitized reed warbler nests with an artificial non-mimetic cuckoo egg, and then monitored the reaction of the hosts. In 27 out of 48 nests (56.3%) the parasitic egg was rejected. The rejecter pairs had a statistically significantly lower intraclutch variation in egg appearance than the acceptor pairs. We discuss possible explanations for the observed relationship between rejection of unlike eggs and intraclutch variation in egg appearance within this population of reed warblers. The results are consistent with the evolutionary arms race hypothesis, but the intermediate rejection rate found in this population could also be maintained by an equilibrium between acceptors and rejecters due to rejection costs.     

Continuous Variation Rather than Specialization in the Egg Phenotypes of Cuckoos (Cuculus canorus) Parasitizing Two Sympatric Reed Warbler Species

The evolution of brood parasitism has long attracted considerable attention among behavioural ecologists, especially in the common cuckoo system. Common cuckoos (Cuculus canorus) are obligatory brood parasites, laying eggs in nests of passerines and specializing on specific host species. Specialized races of cuckoos are genetically distinct. Often in a given area, cuckoos encounter multiple hosts showing substantial variation in egg morphology. Exploiting different hosts should lead to egg-phenotype specialization in cuckoos to match egg phenotypes of the hosts. Here we test this assumption using a wild population of two sympatrically occurring host species: the great reed warbler (Acrocephalus arundinaceus) and reed warbler (A. scirpaceus). Using colour spectrophotometry, egg shell dynamometry and egg size measurements, we studied egg morphologies of cuckoos parasitizing these two hosts. In spite of observing clear differences between host egg phenotypes, we found no clear differences in cuckoo egg morphologies. Interestingly, although chromatically cuckoo eggs were more similar to reed warbler eggs, after taking into account achromatic differences, cuckoo eggs seemed to be equally similar to both host species. We hypothesize that such pattern may represent an initial stage of an averaging strategy of cuckoos, that – instead of specializing for specific hosts or exploiting only one host – adapt to multiple hosts.     

Parentage and host preference in the common cuckoo Cuculus canorus

Microsatellite DNA markers were used to investigate parentage relationships in a population of common cuckoo Cuculus canorus. Thirty adults and 55 nestlings were genotyped at six loci from blood samples collected over a four‐year period. To test whether each cuckoo female specialises in parasitising one single host species (Host Preference Hypothesis), the maternal relationships were used to record each female's host choice. The results supported the Host Preference Hypothesis since no female (N=3) was recorded to have parasitised more than one of four congeneric host species breeding in the area. In contrast, the males (N=4) did not show such specialisation since two of them sired offspring reared by different host species.     

Parentage and host preference in the common cuckoo Cuculus canorus

Microsatellite DNA markers were used to investigate parentage relationships in a population of common cuckoo Cuculus canorus . Thirty adults and 55 nestlings were genotyped at six loci from blood samples collected over a four-year period. To test whether each cuckoo female specialises in parasitising one single host species (Host Preference Hypothesis), the maternal relationships were used to record each female's host choice. The results supported the Host Preference Hypothesis since no female (N=3) was recorded to have parasitised more than one of four congeneric host species breeding in the area. In contrast, the males (N=4) did not show such specialisation since two of them sired offspring reared by different host species.