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.     

Cuckoos versus reed warblers: Adaptations and counteradaptations

At study sites in Cambridgeshire, England, the percentage of reed warbler, Acrocephalus scirpaceus, nests parasitized by cuckoos, Cuculus canorus, in 2 years was 22·5% and 9·1%. The warblers rejected cuckoo eggs at 19% of parasitized nests. Parasitized clutches suffered less predation than unparasitized clutches, suggesting that the cuckoo itself was the major predator, plundering nests too advanced for parasitism so that the hosts would re-lay. The cuckoos laid a mimetic egg, parasitized nests in the afternoons during the host laying period, usually removed one host egg, laid a remarkably small egg and laid very quickly. Nests were experimentally parasitized with model eggs to study the significance of this procedure. Experiments showed that host discrimination selects for: (1) egg mimicry by cuckoos (poorer matching model eggs were more likely to be rejected); (2) parasitism during the laying period (mimetic eggs put in nests before host laying began were rejected); (3) afternoon laying (mimetic eggs were less likely to be accepted in the early morning than in the afternoon, when hosts were more often absent from the nest); (4) a small egg (large eggs, typical of non-parasitic cuckoos, were more likely to be rejected); (5) rapid laying (a stuffed cuckoo on the nest stimulated increased rejection of model eggs), and (6) sets a limit to host egg removal by cuckoos (if more than one or two are removed desertion may occur). Mimicry may also be selected for because it reduced the chance that second cuckoos can discriminate the first cuckoo's egg from the host's clutch. Predation did not select for mimicry; nests with a non-mimetic egg did not suffer greater predation than those with a mimetic egg. Host rejection of model eggs did not depend on: (1) stage of parasitism once host egg laying had begun (nevertheless cuckoos were more likely to lay early in the host laying period probably to increase the chance the cuckoo chick hatched); (2) removal of a host egg (however, this reduced the incidence of unhatched eggs so cuckoos may remove a host egg so as not to exceed the host incubation limit). There were two costs of rejection, an ‘ejection’ cost (own eggs ejected as well as the cuckoo egg) and, with mimetic eggs, a ‘recognition’ cost (own eggs ejected instead of the cuckoo egg). Reed warblers did not discriminate against unlike chicks (another species) and did not favour either a cuckoo chick or their own chicks when these were placed in two nests side by side. Possible reasons why the hosts discriminate against unlike eggs but not unlike chicks are discussed.     

The comparative breeding behaviour of two sympatric cuckoos, Horsfield's Bronze-Cuckoo Chrysococcyx basalts and the Shining Bronze-Cuckoo C. lucidus, in Western Australia: a new model for the evolution of egg morphology and host specificity in avian brood parasites

The breeding behaviour of two similarly sized sympatric cuckoos, Horsfield's Bronze-Cuckoo Chrysococcyx basalts and the Shining Bronze-Cuckoo C. lucidus, was studied over four breeding seasons at Gooseberry Hill, Western Australia. Both cuckoos usually began laying in late August; Shining Bronze-Cuckoos laid for up to 13 weeks and Horsfield's Bronze-Cuckoos for up to 15 weeks. Four host species were parasitized and major hosts were parasitized throughout most of their laying periods. The frequency of parasitism varied between hosts and between years, but Splendid Fairy-wrens Malurus splendens and Yellow-rumped Thornbills Acanthiza chrysorrhoa (major hosts) were always parasitized more heavily than Western Thornbills A. inornata and Scarlet Robins Petroica multicolor. Western Thornbills were parasitized by both cuckoos. Horsfield's and Shining Bronze-Cuckoos laid monomorphic eggs; those of Horsfield's Bronze-Cuckoos were highly mimetic whereas those of Shining Bronze-Cuckoos were non-mimetic and dark in colour. Both cuckoos laid one egg per host nest, deposited eggs directly into the nest, laid very quickly in the early morning, removed at least one host egg at laying, laid eggs small for the size of the birds, hatched after 12 days and evicted nest companions shortly after hatching. Laying was well synchronized with the start of incubation by hosts. Field observations and experiments with egg models indicated that neither of the major hosts, nor the secondary host in common, discriminate against foreign eggs. The nestling period for Horsfield's Bronze-Cuckoo was 17 days, and for the Shining Bronze-Cuckoo 20 days. There was a corresponding difference in nestling growth rate between the cuckoo species. About 50% of cuckoo eggs produced fledglings. Reproductive success for both cuckoos was highest in nests of the secondary host in common, the Western Thornbill. Young cuckoos reached independence 5–6 weeks after hatching. The adaptive significance of competition between cuckoos as a selective agent for cuckoo egg morphology and host specificity is discussed.     

Cuckoo females preferentially use specific habitats when searching for host nests

Egg mimicry is an important adaptation of common cuckoos, Cuculus canorus, against rejection of eggs by their respective hosts. A precondition for the maintenance of egg mimicry is that female cuckoos find hosts with a matching egg type. Experimental evidence indicated that habitat imprinting may be important for host selection. We tested whether the spacing and laying patterns of female cuckoos in the field are compatible with the supposed habitat-imprinting mechanism. We observed 16 females, with the help of radiotelemetry; of seven females, we observed directly 26 egg layings and 27 nest visits without laying. As expected if females were imprinted on different vegetation types, (1) the distribution of vegetation types differed between female home ranges, (2) female habitat use differed from average habitat availability within the egg-laying area (habitat preference), (3) females visited nests and deposited their eggs in the habitat they preferred, and (4) females laid their eggs consistently in a particular habitat type, irrespective of the host species. These results indicate that cuckoo females show habitat preference when searching for suitable host nests. Hence our data are compatible with the habitat-imprinting hypothesis, but owing to the habitat specificity of hosts the data cannot disprove a potential role of host specificity in cuckoo females.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Cryptic cuckoo eggs hide from competing cuckoos

Interspecific arms races between cuckoos and their hosts have produced remarkable examples of mimicry, with parasite eggs evolving to match host egg appearance and so evade removal by hosts. Certain bronze-cuckoo species, however, lay eggs that are cryptic rather than mimetic. These eggs are coated in a low luminance pigment that camouflages them within the dark interiors of hosts'' nests. We investigated whether cuckoo egg crypsis is likely to have arisen from the same coevolutionary processes known to favour egg mimicry. We added high and low luminance-painted eggs to the nests of large-billed gerygones (Gerygone magnirostris), a host of the little bronze-cuckoo (Chalcites minutillus). Gerygones rarely rejected either egg type, and did not reject natural cuckoo eggs. Cuckoos, by contrast, regularly removed an egg from clutches before laying their own and were five times more likely to remove a high luminance model than its low luminance counterpart. Given that we found one-third of all parasitized nests were exploited by multiple cuckoos, our results suggest that competition between cuckoos has been the key selective agent for egg crypsis. In such intraspecific arms races, crypsis may be favoured over mimicry because it can reduce the risk of egg removal to levels below chance.     

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.     

Egg shape mimicry in parasitic cuckoos

Parasitic cuckoos lay their eggs in nests of host species. Rejection of cuckoo eggs by hosts has led to the evolution of egg mimicry by cuckoos, whereby their eggs mimic the colour and pattern of their host eggs to avoid egg recognition and rejection. There is also evidence of mimicry in egg size in some cuckoo–host systems, but currently it is unknown whether cuckoos can also mimic the egg shape of their hosts. In this study, we test whether there is evidence of mimicry in egg form (shape and size) in three species of Australian cuckoos: the fan‐tailed cuckoo Cacomantis flabelliformis, which exploits dome nesting hosts, the brush cuckoo Cacomantis variolosus, which exploits both dome and cup nesting hosts, and the pallid cuckoo Cuculus pallidus, which exploits cup nesting hosts. We found evidence of size mimicry and, for the first time, evidence of egg shape mimicry in two Australian cuckoo species (pallid cuckoo and brush cuckoo). Moreover, cuckoo–host egg similarity was higher for hosts with open nests than for hosts with closed nests. This finding fits well with theory, as it has been suggested that hosts with closed nests have more difficulty recognizing parasitic eggs than open nests, have lower rejection rates and thus exert lower selection for mimicry in cuckoos. This is the first evidence of mimicry in egg shape in a cuckoo–host system, suggesting that mimicry at different levels (size, shape, colour pattern) is evolving in concert. We also confirm the existence of egg size mimicry in cuckoo–host systems.     

Breeding success of common cuckoos Cuculus canorus parasitising four sympatric species of Acrocephalus warblers

We investigated the level of parasitism, egg mimicry and breeding success of cuckoos parasitising four sympatric species of Acrocephalus warblers in southern Moravia, Czech Republic. The parasitism rate was highest in the marsh warbler Acrocephalus palustris (44.8%) followed by great reed warbler A. arundinaceus (33.8%), sedge warbler A. schoenobaenus (26.5%) and reed warbler A. scirpaceus (11.6%). Although the cuckoo eggs showed a high level of mimicry the eggs of the marsh warbler this host species rejected 72% of the cuckoo eggs, resulting in a cuckoo breeding success of only 4.3%. Cuckoo eggs laid in great reed warbler and reed warbler nests showed a similar hatching success, but the cuckoo chicks survived better in great reed warbler nests, resulting in a breeding success of 30.4%, as compared to 16.4% in nests of the reed warbler. The relationship between the level of parasitism, host rejection of cuckoo eggs, cuckoo chick survival and breeding success is discussed for the four host species.     

THE OCCURRENCE AND SIGNIFICANCE OF ANOMALOUS REPRODUCTIVE ACTIVITIES IN TWO NORTH AMERICAN NON-PARASITIC CUCKOOS COCCYZUS SPP.

Among the unusual breeding habits of the non-parasitic Yellow-billed and Black-billed Cuckoos of North America are great variability in clutch size and rate of laying, initiation of incubation long before the clutch is complete, occasional laying in nests of other species, annual irregularity in the timing of the breeding season, and semi-nomadic post-migratory movements into breeding areas where food is abundant. These facts, in addition to their peculiar diet and the very large size of their eggs, suggest that cuckoos have extraordinary problems in obtaining adequate energy for reproduction. At Bloomington, Indiana (U.S.A.), during a 15-year period, anomalies in the reproductive activities of cuckoos were concentrated into two years in which food was abundant. This was particularly true of one of these years, when there was a vast emergence of periodical cicadas: the Yellow-billed Cuckoo advanced its normal schedule and bred during peak cicada abundance, laid unusually large clutches, and parasitized Black-billed Cuckoo nests. Some females may have resumed laying in nests in which, having already deposited clutches of normal size, they had been incubating for long periods; the alternative possibility is that there was intraspecific brood parasitism. The erratic egg-laying behaviour of these cuckoos is attributed to the evolution of mechanisms permitting very quick exploitation of a favourable feeding situation. It is suggested that reproductive behaviour has become so responsive to an abundance of food that normal ordering and integration of the stages of breeding have been lost in some females. Such a loss could be responsible for the laying of eggs in alien nests, and it may have been the antecedent of obligate brood parasitism in parasitic cuckoo species.     

Habitat-dependent call divergence in the common cuckoo: is it a potential signal for assortative mating?

The common cuckoo (Cuculus canorus) is an obligate brood parasite that mimics the eggs of its hosts. The host-specific egg pattern is thought to be inherited matrilinearly, creating female-only host-specific races. Males are thought not to be adapted to their host and they maintain the species by mating arbitrarily with respect to host specialization of females. However, recent results suggest that male cuckoos may also show host-specific adaptations and these may require assortative mating with respect to host. The calls males produce on the breeding grounds could provide a potential mechanism for assortative mating. We tested whether male cuckoo calls differ more between nearby populations that parasitize different hosts than between distant populations that parasitize the same host. We recorded the calls of geographically distant pairs of populations in Hungary, with each pair consisting of a forest population and a nearby reed bed population. Each habitat is characterized by one main host species for the common cuckoo. Our results show that calls of distant cuckoo populations from the same habitat type are more similar to each other than they are to those of nearby populations from a different habitat. These results suggest that cuckoo calls differ sufficiently to allow recognition of habitat-specific individuals.     

Rapid increase in cuckoo egg matching in a recently parasitized reed warbler population

Parasitic cuckoos lay eggs that mimic those of their hosts, and such close phenotypic matching may arise from coevolutionary interactions between parasite and host. However, cuckoos may also explicitly choose hosts in a way that increases degree of matching between eggs of cuckoos and parasites, with female preference for specific host phenotypes increasing the degree of matching. We tested for temporal change in degree of matching between eggs of the parasitic European cuckoo (Cuculus canorus) and its reed warbler (Acrocephalus scirpaceus) host during 24 consecutive years in a recently parasitized reed warbler population. Cuckoo-host egg matching in an ultraviolet-brownness component yielding most of the chromatic variance of eggs improved during the study period. Improved matching was not due to changes in cuckoo egg phenotype. Cuckoo eggs matched host eggs for ultraviolet-brownness within nests irrespective of duration of sympatry. Ultraviolet-brownness of cuckoo eggs was similar to that of reed warbler eggs at parasitized nests, but differed from that of reed warbler eggs at unparasitized nests. These findings provide tentative support for the cuckoo preference hypothesis suggesting that cuckoo-host egg matching could partially be due to cuckoo females selecting host nests based on the appearance of their eggs.     

Do cuckoos choose nests of great reed warblers on the basis of host egg appearance?

Prevailing theory assumes cuckoos lay at random among host nests within a population, although it has been suggested that cuckoos could choose large nests and relatively active pairs within host populations. We tested the hypothesis that egg matching could be improved by cuckoos choosing nests in which host eggs more closely match their own, by assessing matching and monitoring nest fate in great reed warblers naturally or experimentally parasitized by eggs of European cuckoos. A positive correlation between cuckoo and host egg visual features suggests that cuckoos do not lay at random within a population, but choose nests and this improves egg matching: naturally parasitized cuckoo eggs were more similar to host eggs as perceived by humans and as measured by spectrophotometry. Our results suggest a hitherto overlooked step in cuckoo-host evolutionary arms races, and have nontrivial implications for the common experimental practice of artificially parasitizing clutches.     

How is host egg mimicry maintained in the cuckoo (Cuculus canorus)?

To investigate the evolutionary mechanism (host specificity vs. random searching) maintaining mimicry between cuckoo egg appearance and that of different European cuckoo Cuculus canorus hosts, we studied the level of mimicry between the appearance of C. canorus eggs and that of their hosts' eggs in different habitats in southern Finland by using ultraviolet-visible reflectance spectrophotometry. In the main habitat used by C. canorus for reproduction, eggs laid in nests of different host species differed in appearance. Host use by C. canorus was not related to the abundance of hosts, and the level of mimicry was not related to host abundance in the habitat. Furthermore, a close match between C. canorus egg appearance and that of host eggs within habitats was detected after removing the potentially confounding effect of host abundance. In the only two suitable host species nesting in trees (namely chaffinch Fringilla coelebs and brambling Fringilla montifringilla ) we detected changes in C. canorus egg appearance that paralleled those of the two host species. Thus our findings suggest the existence of a correlation between the appearance of C. canorus eggs and that of their hosts' eggs within different habitat types, and suggest that mimicry is maintained by strict host preferences by each C. canorus female when laying.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 57–68.     

Do common cuckoos (Cuculus canorus) possess an optimal laying behaviour to match their own egg phenotype to that of their Oriental reed warbler (Acrocephalus orientalis) hosts?

Optimality theory suggests that parasitic cuckoos should evolve an optimal laying behaviour aiming to positively select host nests in which the eggs match the phenotype of their own eggs, thus minimizing the rejection risk from hosts and, in turn, maximizing the cuckoos' fitness. We tested this hypothesis by investigating cuckoo‐egg matching between parasitized and nonparasitized nests in a common cuckoo (Cuculus canorus) host, the Oriental reed warbler (Acrocephalus orientalis), by use of Vorobyev–Osorio and Nature‐Pattern‐Match models to quantify the matching of egg colour and pattern from avian vision, respectively. The results of our study indicated that cuckoo‐egg matching in parasitized nests was no better than that in nonparasitized nests, and thus we found no support for the optimal laying hypothesis in cuckoos. The mixed conclusions from all previous studies, including the present study, may be explained by (1) the parallel coevolution in different cuckoo–host systems; (2) the inappropriate methodology; and (3) the deficiency of the assumption itself. We suggest that a better methodology should be developed to solve the puzzle of whether cuckoos choose to lay eggs matching those of the host.     

Absence of egg discrimination in a suitable cuckoo Cuculus canorus host breeding away from trees

There is at present considerable variation in the level of antiparasite defences among different host species of avian brood parasites, but in many potential hosts some individuals reject poorly matching parasite eggs. Here we present unique absence of egg discrimination behaviour backed up by a lack of egg recognition abilities in a suitable common cuckoo Cuculus canorus host, the skylark Alauda arvensis. Skylarks did not show any clear rejection response to experimentally added highly non‐mimetic foreign eggs in any behavioural context, even before they had started laying or when the whole clutch was exchanged with foreign eggs. This absence of antiparasite defence can be explained by the breeding habitat of larks consisting of largely treeless open landscapes where cuckoos have little access to the nests, thereby eroding the possibility of coevolutionary interactions. Our results are strikingly consistent with the spatial habitat structure hypothesis proposed to explain the occurrence and extent of avian host‐parasite co‐adaptation.     

Cuckoo parasitism in a cavity nesting host: near absent egg‐rejection in a northern redstart population under heavy apparent (but low effective) brood parasitism

Brood parasite – host systems continue to offer insights into species coevolution. A notable system is the redstart Phoenicurus phoenicurus parasitized by the ‘redstart‐cuckoo’ Cuculus canorus gens. Redstarts are the only regular cuckoo hosts that breed in cavities, which challenges adult cuckoos in egg laying and cuckoo chicks in host eviction. We investigated parasitism in this system and found high overall parasitism rates (31.1% of 360 redstart nests), but also that only 33.1% of parasitism events (49 of 148 eggs) were successful in laying eggs into redstart nest cups. The majority of cuckoo eggs were mislaid and found on the rim of the nest; outside the nest cup. All available evidence suggests these eggs were not ejected by hosts. The effective parasitism rate was therefore only 12.8% of redstart nests. Redstarts responded to natural parasitism by deserting their nests in 13.0% of cases, compared to desertion rates of 2.8% for non‐parasitized nests. Our egg parasitism experiments found low rates (12.2%) of rejection of artificial non‐mimetic cuckoo eggs. Artificial mimetic and real cuckoo eggs added to nests were rejected at even lower rates, and were always rejected via desertion. Under natural conditions, only 21 cuckoo chicks fledged of 150 cuckoo eggs laid. Adding to this low success, is that cuckoo chicks are sometimes unable to evict all host young, and were more likely to die as a result compared to cuckoo chicks reared alone. This low success seems to be mainly due to the cavity nesting strategy of the redstart which is a challenging obstacle for the cuckoo. The redstart‐cuckoo system appears to be a fruitful model system and we suggest much more emphasis should be placed on frontline defences such as nest site selection strategies when investigating brood parasite–host coevolution.     

Striking difference in response to expanding brood parasites by birds in western and eastern Beringia

Two species of obligate brood‐parasitic Cuculus cuckoos are expanding their ranges in Beringia. Both now breed on the Asian side, close to the Bering Strait, and are found in Alaska during the breeding season. From May to July 2017, we used painted 3D‐printed model eggs of two cuckoo host‐races breeding in northeastern Siberia to test behavioral responses of native songbirds on both sides of the Bering Strait, with particular attention to species that are known cuckoo hosts in their Siberian range. Each host nest was tested after the second egg was laid and, if possible, again 4 days later with a model of a different type. Although our Siberian study site was also outside the known breeding ranges of the cuckoos, we found that Siberian birds had strong anti‐parasite responses, with 14 of 22 models rejected. In contrast, birds in Alaska had virtually no detectable anti‐parasite behaviors, with only one of 96 models rejected; the rejecters were Red‐throated Pipits (Anthus cervinus). Such differences suggest that the cuckoos might successfully parasitize naïve hosts and become established in North America whether or not their historic host species are widely available.     

Cryptic gentes revealed in pallid cuckoos Cuculus pallidus using reflectance spectrophotometry

Many cuckoo species lay eggs that match those of their hosts, which can significantly reduce rejection of their eggs by the host species. However, egg mimicry is problematic for generalist cuckoos that parasitize several host species with different egg types. Some generalist cuckoos have overcome this problem by evolving several host-specific races (gentes), each with its own, host-specific egg type. It is unknown how generalist cuckoos lacking gentes are able to avoid egg rejection by hosts. Here we use reflectance spectrophotometry (300-700 nm) on museum egg collections to test for host-specific egg types in an Australian generalist cuckoo reported to have a single egg type. We show that the colour of pallid cuckoo (Cuculus pallidus) eggs differed between four host species, and that their eggs closely mimicked the eggs of the host they parasitized. These results reveal that pallid cuckoos have host-specific egg types that have not been detected by human observation, and indicate that gentes could be more common than previously realized.     

Explaining variation in brood parasitism rates between potential host species with similar habitat requirements

Host specialization evolved in many parasite-host systems. Evolution and maintenance of host specificity may be influenced by host life-history traits, active host selection by the parasite, and host anti-parasite strategies. The relative importance of these factors is poorly understood in situations that offer parasites a choice between hosts with similar habitat requirements. The common cuckoo Cuculus canorus is a generalist parasite on the species level, but individual females prefer particular host species. In reed beds of the Yellow River Delta, China, two potential hosts with similar nest characteristics, Oriental reed warblers Acrocephalus orientalis and reed parrotbills Paradoxornis heudei, breed in sympatry. We found that warblers were parasitized at much higher rates than parrotbills. Both hosts recognized and rejected non-mimetic model eggs well, indicating that they have been involved in an arms-race with cuckoos. Cuckoo eggs closely resembled warbler eggs, and such eggs were mostly accepted by warblers but rejected by parrotbills. Only warblers recognized adult cuckoos as a specific threat. Both hosts were equally good at raising cuckoo chicks. Low nest density, partial isolation by breeding time, small scale differences in nest and nest site characteristics, and high rejection rates of natural cuckoo eggs are likely cumulatively responsible for the low current parasitism rate in parrotbills. This study emphasizes the importance of integrating the study of general host life-history characteristics and specific anti-parasitism strategies of hosts across all breeding stages to understand the evolution of host specificity.     

A comparative study of host selection in the European cuckoo Cuculus canorus

Certain kinds of hosts are commonly regarded as being more suitable than other for rearing European cuckoos (Cuculus canorus) – insectivores that lay small eggs and have open, shallow nests – although empirical tests of cuckoo host selection are lacking. We analysed host use by the European cuckoo in 72 British passerines that are potential hosts and for which there was information available on life-history variables and variables related to cuckoo-host coevolution, such as rate of parasitism, rejection rate of non-mimetic model eggs and degree of cuckoo-egg mimicry of host eggs. The relative population size of the host species affected parasitism rate most strongly, followed by relatively short duration of the nestling period, and the kind of nest, with cuckoos selecting open-nesting hosts. However, the effect of the nestling period could be related to host body size and the kind of nest used, because hole-nesting species normally have longer nestling periods than open-nesters. We re-analysed the data excluding hole nesters and corvid species (species with larger body mass), but the results remained identical. The European cuckoo may benefit from selecting hosts with short nestling periods because such hosts provide food for their nestlings at a very high rate. When only those species known as cuckoo hosts were analysed, the variable that best accounted for the parasitism rate was duration of the breeding season. Therefore, availability of potential hosts in both time and space is important for cuckoos in selecting hosts. Received: 16 July 1998 / Accepted: 27 October 1998