Modelling the maintenance of egg polymorphism in avian brood parasites and their hosts.

In avian brood parasitism, egg phenotype plays a key role for both host and parasite reproduction. Several parrotbill species of the genus Paradoxornis are parasitized by the common cuckoo Cuculus canorus, and clear polymorphism in egg phenotype is observed. In this article, we develop a population genetics model in order to identify the key parameters that control the maintenance of egg polymorphism. The model analyses show that egg polymorphism can be maintained either statically as an equilibrium or dynamically with frequency oscillations depending on the sensitivity of the host against unlike eggs and how the parasite targets host nests with specific egg phenotypes. On the basis of the model, we discuss egg polymorphism observed in parrotbills and other host species parasitized by the cuckoo. We suggest the possibility that frequencies of egg phenotypes oscillate and we appeal for monitoring of cuckoo-host interactions over a large spatiotemporal scale.

Rapid change in host use of the common cuckoo Cuculus canorus linked to climate change.

Parasites require synchrony with their hosts so if host timing changes with climate change, some parasites may decline and eventually go extinct. Residents and short-distance migrant hosts of the brood parasitic common cuckoo, Cuculus canorus, have advanced their phenology in response to climate change more than long-distance migrants, including the cuckoo itself. Because different parts of Europe show different degrees of climate change, we predicted that use of residents or short-distance migrants as hosts should have declined in areas with greater increase in spring temperature. Comparing relative frequency of parasitism of the two host categories in 23 European countries before and after 1990, when spring temperatures in many areas had started to increase, we found that relative parasitism of residents and short-distance migrants decreased. This change in host use was positively related to increase in spring temperature, consistent with the prediction that relative change in phenology for different migrant classes drives host-use patterns. These findings are consistent with the hypothesis that climate change affects the relative abundance of different host races of the common cuckoo.

Isolation by time and habitat and coexistence of distinct host races of the common cuckoo.

Isolation by time occurs when different populations of a single species reproduce at different times and thereby reduce the probability of interbreeding, potentially causing divergent adaptation to timing of reproduction, eventually resulting in ecological species separated by timing of reproduction. We analysed extensive data on timing of reproduction by different host races of the common cuckoo Cuculus canorus that is an obligate brood parasite laying eggs in the nests of many different species of passerine birds. Because different hosts breed at different times, specific host races of cuckoos have adapted to specific hosts by laying eggs when nests of these hosts are available, and such divergence may be further exaggerated by differences in timing of breeding among host races with similar habitat requirements. Host species accounted for a quarter of the variance in timing of breeding by the cuckoo. Common cuckoos reproduced at a similar, but narrower subset of dates as did possible hosts, showing that only a fraction of hosts with specific breeding dates were parasitized. Common cuckoo eggs laid in the 'right' kind of nests, phenotypically matching the eggs of the host, were laid later during the season than cuckoo eggs laid in the 'wrong' kind of nests where the eggs did not mimic those of the host. Pairs of sympatric cuckoo host races differed more in timing of breeding than pairs of allopatric host races, and pairs of cuckoo host races with similar breeding habitat differed more in breeding date than pairs of cuckoo host races with dissimilar habitat, as expected from reproductive character displacement. These findings are consistent with cuckoo host races being isolated by timing of breeding and habitat.

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.

Avian colour perception predicts behavioural responses to experimental brood parasitism in chaffinches.

Hosts of cuckoos have evolved defences allowing them to discriminate and reject parasite eggs. Mechanisms of discrimination are mostly visually mediated, and have been studied using approaches that do not account for what the receiver (i.e. host) actually can discriminate. Here, for the first time we apply a perceptual model of colour discrimination to study behavioural responses to natural variation in parasite egg appearance in chaffinches Fringilla coelebs. Discrimination of parasite eggs gradually increased with increasing differences in chromatic contrasts as perceived by birds between parasite and host eggs. These results confirm that colour differences of the eggs as perceived by birds are important integral parts of a matching signal used by chaffinch hosts.

Egg phenotype differentiation in sympatric cuckoo Cuculus canorus gentes.

The brood parasitic common cuckoo Cuculus canorus consists of gentes, which typically parasitize only a single host species whose eggs they often mimic. Where multiple cuckoo gentes co-exist in sympatry, we may expect variable but generally poorer mimicry because of host switches or inter-gens gene flow via males if these also contribute to egg phenotypes. Here, we investigated egg trait differentiation and mimicry in three cuckoo gentes parasitizing great reed warblers Acrocephalus arundinaceus, marsh warblers Acrocephalus palustris and corn buntings Miliaria calandra breeding in close sympatry in partially overlapping habitat types. The three cuckoo gentes showed a remarkable degree of mimicry to their three host species in some but not all egg features, including egg size, a hitherto largely ignored feature of egg mimicry. Egg phenotype matching for both background and spot colours as well as for egg size has been maintained in close sympatry despite the possibility for gene flow.

Cell-mediated immunity and multi-locus heterozygosity in bluethroat nestlings.

Recent evidence suggests that marker-based heterozygosity-fitness correlations may be driven by only one or a few markers, indicating local heterozygosity effects caused by linkage disequilibrium with functional genes. In this study, we investigated the relationship between microsatellite heterozygosity and a measure of cell-mediated immunity (phytohaemagglutinin; PHA) in bluethroat (Luscinia s. svecica) nestlings using a full-sibling design. We found significant positive associations between PHA response and two different indices of microsatellite heterozygosity, i.e. multi-locus heterozygosity and mean d(2). However, model comparisons disclosed that both associations were more likely caused by local effects rather than general effects and that the two local effects appeared to be realized through two different genetic mechanisms. Our results indicate that both the random assortment of parental chromosomes during meiosis as well as inbreeding can drive heterozygosity-fitness correlations.

Accumulation of heavy metals in circumpolar willow ptarmigan (Lagopus l. lagopus) populations.

A circumpolar survey of heavy metals in willow ptarmigan liver and kidney revealed considerable variations in Cd content in Canada and Scandinavia. The Cd content in central Canada was comparable with that in Scandinavia and Russia, at least for kidney. However, in both liver and kidney the median for Canada as a whole was much higher than in the other countries. Some Canadian locations had exceptionally high levels, several birds having >50 mg kg(-1) in liver and >400 mg kg(-1) in kidney. In Norway, the Cd content was highest in central mountain areas in south Norway and inland locations in the two northernmost counties. Five locations in central and north Norway showed mean Cd levels in kidney above 100 mg kg(-1). It is difficult to evaluate regional differences in Sweden, but most locations had the same Cd level as moderately contaminated locations in Norway. Cd levels in Russia were comparable to moderately contaminated locations in the other countries. Due to a high intake of willow, naturally rich in Cd, direct comparison of the Cd level in willow ptarmigan from different locations cannot reveal the effects of long-range pollution. The Pb concentration in willow ptarmigan kidney and liver varied significantly in Norway and Canada, but not in Sweden and Russia. Levels in Sweden and Russia were comparable to those in Canada and low levels in Norway. The highest median value from all locations within countries was found in Norway, both in liver and kidney. The highest Pb content was found in south Norway, indicating an effect of long-range pollution in willow ptarmigan. The level in western Canada was significantly higher than in central Canada. The Hg content in liver varied significantly from one location to another in all the countries and in kidney everywhere except Sweden. In Scandinavia, there is no distinct regional pattern. Canada had a significantly higher Hg level in central than western regions in both tissues the opposite of that found for Cd and Pb. Cu and Zn showed significant variations from one location to another in liver and kidney in Canada and Norway, but only in kidney in samples from Sweden. Comparison between western and central Canada revealed a significant difference for Cu in liver, samples from central Canada having more. There are no significant differences from one country to another, but some localities in Canada seem to have higher Cu concentrations in kidney than are found in Scandinavia and Russia.