Importance of communal foraging grounds outside the reed marsh for breeding great reed warblers

Foraging habitat selection of breeding great reed warblers was studied at a shore of Lake Biwa. The foraging grounds of parent warblers during the nesting period were not restricted to the breeding territory of the reed marsh, their nestling habitat. The paddy field outside the reed marsh was used communally by them throughout the breeding season. Females with early stage nestlings did not visit the paddy field whereas when nestlings were older than 3 days, more than half of their total food was collected there. Females with nests adjacent to the paddy field tended to exploit the paddy field more often than those with nests distant from it. Monogamously mated females tended to exploit the paddy field more often than polygynously mated females. Food collected in the paddy field was larger than that in the reed marsh and parent birds were prepared to travel longer distances to exploit the rich source of food in the paddy field. The importance of the communal foraging ground outside the reed marsh as a background of the polygynous mating system of this species is discussed.     

Hybridization but no evidence for backcrossing and introgression in a sympatric population of great reed warblers and clamorous reed warblers

Hybridization is observed frequently in birds, but often it is not known whether the hybrids are fertile and if backcrossing occurs. The breeding ranges of the great reed warbler (Acrocephalus arundinaceus) and the clamorous reed warbler (A. stentoreus) overlap in southern Kazakhstan and a previous study has documented hybridization in a sympatric population. In the present study, we first present a large set of novel microsatellite loci isolated and characterised in great reed warblers. Secondly, we evaluate whether hybridization in the sympatric breeding population has been followed by backcrossing and introgression.We isolated 181 unique microsatellite loci in great reed warblers. Of 41 loci evaluated, 40 amplified and 30 were polymorphic. Bayesian clustering analyses based on genotype data from 23 autosomal loci recognised two well-defined genetic clusters corresponding to the two species. Individuals clustered to a very high extent to either of these clusters (admixture proportions ≥ 0.984) with the exception of four previously suggested arundinaceus-stentoreus hybrid birds that showed mixed ancestry (admixture proportions 0.495-0.619). Analyses of simulated hybrids and backcrossed individuals showed that the sampled birds do not correspond to first-fourth-generation backcrosses, and that fifth or higher generation backcrosses to a high extent resemble 'pure' birds at this set of markers.We conclude that these novel microsatellite loci provide a useful molecular resource for Acrocephalus warblers. The time to reach reproductive isolation is believed to be very long in birds, approximately 5 Myrs, and with an estimated divergence time of 2 Myrs between these warblers, some backcrossing and introgression could have been expected. However, there was no evidence for backcrossing and introgression suggesting that hybrids are either infertile or their progeny inviable. Very low levels of introgression cannot be excluded, which still may be an important factor as a source of new genetic variation.     

Microsatellite diversity predicts recruitment of sibling great reed warblers.

Inbreeding increases the level of homozygosity, which in turn might depress fitness. In addition, individuals having the same inbreeding coefficient (e.g. siblings) vary in homozygosity. The potential fitness effects of variation in homozygosity that is unrelated to the inbreeding coefficient have seldom been examined. Here, we present evidence from wild birds that genetic variation at five microsatellite loci predicts the recruitment success of siblings. Dyads of full-sibling great reed warblers (Acrocephalus arundinaceus), one individual of which became a recruit to the natal population while the other did not return, were selected for the analysis. Each dyad was matched for sex and size. Local recruitment is strongly tied to fitness in great reed warblers as the majority of offspring die before adulthood, philopatry predominates among surviving individuals and emigrants have lower lifetime fitness. Paired tests showed that recruited individuals had higher individual heterozygosity and higher genetic diversity, which was measured as the mean squared distance between microsatellite alleles (mean d(2)), than their non-recruited siblings. These relationships suggest that the microsatellite markers, which are generally assumed to be neutral, cosegregated with genes exhibiting genetic variation for fitness.     

Daily energy expenditure of singing great reed warblers Acrocephalus arundinaceus

According to honest signalling theory, signals must be costly to produce to retain information about the signaller's quality. The song produced by male birds during breeding is a vocal "ornament" used for intra- and inter-sexual purposes. The energetic cost of this vocal signal remains a contentious issue. We used the doubly labelled water method to measure field metabolic rate by estimating CO₂ production and then convert this to daily energy expenditure (DEE) in great reed warbler males singing under natural conditions (10 at low to moderate intensity and 7 at very high intensity from dawn to dusk). There was a significant positive relationship between singing intensity and DEE. From this relationship we extrapolated the average DEE for intensely singing males (i.e., males producing song sounds 50% of the time and hence sitting at their elevated song post in the top of a reed stem more or less continuously throughout the ∼20 h of daylight) to 3.3×BMR (basal metabolic rate) and for non-singing males to 2.2×BMR. The mean DEE measured for the seven males singing with very high intensity was 3.1×BMR. The maximum measured DEE for a single male was 3.9×BMR, i.e. close to the maximum sustainable DEE (4×BMR), and the minimum DEE was 2.1×BMR for a male singing at very low intensity. These results imply that producing intensive advertising song in birds may incur a substantial cost in terms of increased energy expenditure.     

Sex ratio variation among broods of great reed warblers Acrocephalus arundinaceus

The sex of 746 great reed warbler fledglings (from 175 broods) was determined by the use of single primer polymerase chain reaction. The reliability of the technique was confirmed as 104 of the fledglings were subsequently recorded as adults of known sex. The overall sex ratio did not differ from unity. Variation in sex ratios between broods was larger than expected from a binomial distribution. Female identity explained some of the variation of brood sex ratio indicating that certain females consistently produced sex ratios that departed from the average value in the population. The theory of sex allocation predicts that parents should adjust the sex ratio of their brood to the relative value of sons and daughters and this may vary in relation to the quality of the parents or to the time of breeding. In the great reed warbler, the proportion of sons was not related to time of breeding, or to any of five female variables. Of five male variables, males with early arrival date tended to produce more daughters. The sex ratio of fledglings that were a result of extra-pair fertilizations did not differ from that of legitimate fledglings. Hence, there is currently no evidence of that female great reed warblers invest in a higher proportion of sons when mated with attractive males.     

The importance of nest cleaning in egg rejection behaviour of great reed warblers Acrocephalus arandinaceas

We tested the importance of nest cleaning in egg rejection behaviour of the great reed warbler Acrocephalus arundinaceus in a highly parasitised population in which about 64% of nests are parasitised by the common cuckoo Cuculus canorus . Three types of objects of the same weight, texture and colour but with different shapes (dummy cuckoo eggs, sticks and disks) were placed into great reed warbler nests. We investigated the response of hosts in two stages of breeding: pre-incubation when the risk of brood parasitism is high, and during incubation when the risk of parasitism is low. The dummy cuckoo eggs were rejected less often than the other objects in both breeding stages, although we did not find any difference in the frequency of rejection between pre-incubation and incubation. We integrate these results into current views on the evolution of host–parasite interactions, and propose a hierarchical concept to understand egg rejection behaviour: (1) hosts reject all non-egg shaped objects as a general cleaning mechanism; (2) adaptations for the hosts' ability to recognise their own eggs allows them to distinguish these eggs from similar objects and parasitic eggs.     

Primary peak and chronic malaria infection levels are correlated in experimentally infected great reed warblers

SUMMARY Malaria parasites often manage to maintain an infection for several months or years in their vertebrate hosts. In humans, rodents and birds, most of the fitness costs associated with malaria infections are in the short initial primary (high parasitaemia) phase of the infection, whereas the chronic phase (low parasitaemia) is more benign to the host. In wild birds, malaria parasites have mainly been studied during the chronic phase of the infection. This is because the initial primary phase of infection is short in duration and infected birds with severe disease symptoms tend to hide in sheltered places and are thus rarely caught and sampled. We therefore wanted to investigate the relationship between the parasitaemia during the primary and chronic phases of the infection using an experimental infection approach. We found a significant positive correlation between parasitaemia in the primary peak and the subsequent chronic phase of infection when we experimentally infected great reed warblers (Acrocephalus arundinaceus) with Plasmodium ashfordi. The reason for this association remains to be understood, but might arise from individual variation in exoerythrocytic parasite reservoirs in hosts, parasite antigenic diversity and/or host genetics. Our results suggest that the chronic phase parasitaemia can be used to qualitatively infer the parasitaemia of the preceding and more severe primary phase, which is a very important finding for studies of avian malaria in wild populations.     

Polygynous great reed warblers Acrocephalus arundinaceus suffer more cuckoo Cuculus canorus parasitism than monogamous pairs

There is increasing evidence that hosts within a population may not be parasitized by common cuckoos Cuculus canorus with equal probability. Such non‐randomness has been documented, for example, for host nest sites and host quality. In this study we demonstrate association between successful cuckoo parasitism and host social mating system. We found that nests of socially polygynous great reed warblers Acrocephalus arundinaceus were more often successfully parasitized than the nests of their monogamous counterparts. We hypothesize that lack of parental assistance provided by polygynous males to their mates during egg laying period and higher nest activity in their territories could contribute to this discrepancy. These data imply that social mating system should be taken into account in future studies of brood parasitism.     

Nocturnal migratory flight initiation in reed warblers Acrocephalus scirpaceus: effect of wind on orientation and timing of migration

We used radio-telemetry to study autumn migratory flight initiation and orientation in relation to wind and air pressure in a nocturnal passerine migrant, the reed warbler Acrocephalus scirpaceus at Falsterbo, southwest Sweden. The majority of the reed warblers departed in the expected migratory direction towards south of southwest, while a low number of the birds took off in reverse directions between north and east. Flight directions at departure correlated with wind directions. These correlations were particularly prominent at higher wind speeds but were absent at wind speeds below 4 m/s. Birds departing in the expected migratory direction compensated completely for wind drift. The reed warblers preferred to depart during nights with tailwinds and when air pressure was increasing suggesting that reed warblers are sensitive to winds and air pressure and select favourable wind conditions for their migratory flights. Since air pressure as well as velocity and direction of the wind are correlated with the passage of cyclones, a combination of these weather variables is presumably important for the birds' decision to migrate and should therefore be considered in optimal migration models.     

Genetic and phenotypic associations in morphological traits: a long term study of great reed warblers Acrocephalus arundinaceus

We present estimates of standardized selection (directional and quadratic) differentials via reproductive success on eight morphological traits in a newly founded long-term study population of great reed warblers Acrocephalus arundinaceus in Sweden. In order to predict the evolutionary response to selection in these traits we present estimates of heritabilities (h²), phenotypic (r_P) and genetic (r_A) correlations among the same traits. We also examined the extent of parental effects in the expression of the phenotypic traits. Overall, the population is subject to low levels of directional selection and higher levels of stabilizing selection. This makes us predict that the population is changing very little if anything, even though h² in many cases are considerable. Midparent-midoffspring h² ranged from 0.14 and 0.94 (mean 0.58) and were significant for seven of eight traits. We found indications of positive maternal effects in tarsus length. Phenotypic correlations between traits ranged from 0.02 to 0.43 (mean 0.15) and showed generally much lower values than the corresponding genetic correlations that ranged between 0.08 and 1.04 (mean 0.46). Overall, the correlation between r_P and r_A was significant, although moderate, but they tended to differ in magnitude, possibly due to overestimation of additive covariance between traits.     

A strong quantitative trait locus for wing length on chromosome 2 in a wild population of great reed warblers

Wing length is a key character for essential behaviours related to bird flight such as migration and foraging. In the present study, we initiate the search for the genes underlying wing length in birds by studying a long-distance migrant, the great reed warbler (Acrocephalus arundinaceus). In this species wing length is an evolutionary interesting trait with pronounced latitudinal gradient and sex-specific selection regimes in local populations. We performed a quantitative trait locus (QTL) scan for wing length in great reed warblers using phenotypic, genotypic, pedigree and linkage map data from our long-term study population in Sweden. We applied the linkage analysis mapping method implemented in GridQTL (a new web-based software) and detected a genome-wide significant QTL for wing length on chromosome 2, to our knowledge, the first detected QTL in wild birds. The QTL extended over 25 cM and accounted for a substantial part (37%) of the phenotypic variance of the trait. A genome scan for tarsus length (a body-size-related trait) did not show any signal, implying that the wing-length QTL on chromosome 2 was not associated with body size. Our results provide a first important step into understanding the genetic architecture of avian wing length, and give opportunities to study the evolutionary dynamics of wing length at the locus level.     

Stabilising selection on wing length in reed warblers Acrocephalus scirpaceus

The size of an animal is of utmost importance for its overall success and each species is thought to have its own optimal size. If this is true, size traits ought to be under stabilising selection unless the animal is living in a highly unstable environment. Wing length is a standard measurement of the size of birds, but up to date there have been few indications of stabilising selection on wing length. In this study we analyse recovery data for reed warblers Acrocephalus scirpaceus (n=771) ringed as juveniles in Sweden from 1981 until 2001. The data showed a significant relationship between juvenile wing length and survival time, with median juvenile wing lengths (66–67 mm) being the most favourable. The estimated stabilising selection differential (C=−0.094) supports that wing length of the reed warblers in our study is under stabilising selection. The reed warbler is a long-distance migratory species and we suggest that opposing selection pressures may act on wing length during different life history stages, and that this might explain the rather broad peak of favourable wing lengths found.     

Territorial spacing and interspecific competition in three species of reed warblers

Summary Ecological and temporal aspects of the occurrence and dispersion patterns in three reed warbler species were examined. This provided insight into their intra-and interspecific territoriality. Seasonal changes in the occurrence of territorial males in a reed bed along a gradient from the shore landward were studied. The results demonstrate that intra- and interspecific spacing depends on interspecific dominance and/or ecological constraints. In this case it was shown that there is competition for common habitats between reed warblers and great reed warblers, whereas the moustached warbler is ecologically segregated. The reasons for these dispersal patterns are discussed. They do not appear to enhance the availability of food resources. It seems more likely that territorial spacing might function to reduce nest predation.     

Common cuckoo Cuculus canorus parasitism,antiparasite defence and gene flow in closely located populations of great reed warblers Acrocephalus arundinaceus

In Hungary an unusually high rate of parasitism on the great reed warbler Acrocephalus arundinaceus by the common cuckoo Cuculus canorus has been maintained for at least the last one hundred years. We evaluated parasitism rate, antiparasite defence and genetic differentiation among Hungarian great reed warblers at three sites located 40–130 km from each other, where hosts suffered from a high (41–68%), moderate (11%), and almost no (<1%) parasitism. We were especially interested in whether the level of antiparasite defence was related to the local parasitism rate, and, if not, to understand why. There was no difference among the three sites in the responses to experimental parasitism by non‐mimetic model cuckoo eggs (rejection rate 71–82%), which can be explained by strong gene flow between populations: there was low level of philopatry and no genetic differentiation in the region. Reproductive success of the host in the heavily parasitised site was about 54% of that in the unparasitised site, indicating that long‐term persistence of host populations in highly exploited areas depends on continuous immigration.     

Responses of great reed warblers Acrocephalus arundinaceus to experimental brood parasitism: the effects of a cuckoo Cuculus canorus dummy and egg mimicry

Egg rejection behaviour towards parasitic eggs was studied in a great reed warbler Acrocephalus arundinaceus population in central Hungary, which was heavily (about 65%) parasitised by the common cuckoo Cuculus canorus . Clutches were experimentally parasitised during the egg-laying period with artificial, moderately mimetic cuckoo eggs or with conspecific eggs that were good mimics of the hosts' eggs. Great reed warblers rejected 76.2% of the artificial cuckoo eggs, mainly by ejection, but accepted most of the conspecific eggs (87.5%). Cuckoo eggs in naturally parasitised clutches were rejected at a lower rate (32%). When, in addition to the egg mimicry experiments, a stuffed cuckoo was placed near the nest, accompanied by the recording of a female cuckoo call, hosts' rejection rate of the artificial cuckoo egg increased from 76% to 96%. The sight of the cuckoo, on the other hand, did not influence host's rejection behaviour when a conspecific egg was used in the experiment. A stuffed collared dove Streptopelia decaocto , accompanied by its call, was used as a control, and did not cause any increased rejection. Great reed warblers were more aggressive towards the cuckoo than to the dove dummy. When the cuckoo eggs in naturally parasitised clutches were exchanged with artificial cuckoo eggs, we observed no increase in the rejection rate. We conclude that great reed warblers in our heavily parasitised population are capable of detecting brood parasitism in their clutch by identifying the parasitic egg. The efficiency of this identification depends mainly on the mimicry of the foreign egg. The sight of the cuckoo at the nest may increase rejection rate by stimulus summation, and this conditional effect is mainly affected by the degree of mimicry of the parasitic egg.     

Nestling growth and song repertoire size in great reed warblers: evidence for song learning as an indicator mechanism in mate choice

Females of many songbird species show a preference for mating with males that have larger song repertoires, but the advantages associated with this preference are uncertain. We tested the hypothesis that song complexity can serve as an indicator of male quality because the development of the brain regions underlying song learning and production occurs when young birds typically face nutritional and other stresses, so that song reflects how well a male fared during post-hatch development. A key prediction of this hypothesis is that variation in nestling condition should correspond to variation in the adult song repertoires of individuals. We used data from a long-term study of the great reed warbler (Acrocephalus arundinaceus) to test this prediction, correlating two measures of nestling development with subsequent repertoire size of males. We found that the length of the innermost primary feather, a standard measure of development, significantly predicted first-year repertoire size. The relationship between repertoire size and body mass was nearly significant, in spite of the large variance inherent in this measure. These data support the idea that song may provide females with information about a male's response to developmental stress, which in turn is expected to correlate with indirect or direct benefits she might receive.     

Linkage mapping of AFLP markers in a wild population of great reed warblers: importance of heterozygosity and number of genotyped individuals

Amplified fragment length polymorphisms (AFLP) are dominant markers frequently used to build linkage maps where heterozygosity could be inferred by a backcross breeding strategy. In the present study, we describe the utilization of an unmanipulated great reed warbler, Acrocephalus arundinaceus pedigree to infer heterozygous genotypes of AFLP markers in order to map these markers to a partial linkage map previously based on microsatellites. In total, 50 of the 83 autosomal AFLPs (60%) and 4 of 5 Z-linked AFLPs (80%) were mapped. For each marker, on average, 88% of the expected number of heterozygote parents was detected. The likelihood of map assignment was to a large extent due to the number and density of microsatellite markers already in the map. The 'parsimonious linkage map', that is the map based on the most parsimonious location of all significantly linked markers, consisted of 21 autosomal linkage groups with 2 to 15 markers and had a total map size of 552 cM in males and 858 cM in females. The Z-chromosome linkage group with 12 markers had a size of 155 cM. The autosomal 'framework linkage map', that is the map based only on markers with an unambiguous position, had a total size of 237 cM in males and 440 cM in females, respectively. The inclusion of AFLPs enlarged the previous map substantially (e.g. the autosomal parsimonious linkage map became 441 cM and 621 cM larger for male and female recombination, respectively). The probability that an AFLP became mapped increased with increasing level of heterozygosity, whereas the probability of mapping into a framework position increased with both heterozygosity and number of genotyped individuals. Our results suggest that AFLP provides a fast and inexpensive means of enlarging genetic maps already composed of markers with high polymorphism, also in wild populations with unmanipulated pedigrees.     

Systematics of the extinct reed warblers Acrocephalus of the Society Islands of eastern Polynesia

In the Society archipelago (French Polynesia), Acrocephalus reed warblers are known only from four islands: Tahiti, Mo'orea, Huahine and Raiatea. All populations are now extinct except on Tahiti. Our knowledge of these birds is based on a small number of specimens preserved in museums, collected mostly during the 19th century. We present here a review of the past and present distribution, habitat and threats to the Society Islands reed warblers, including details on the specimens in museum collections. We compare the external morphology of the different populations, and use samples from museum specimens to propose a molecular phylogeny of all taxa based on partial cytochrome b gene sequences. The genetic data do not support the monophyly of the Society Islands reed warblers, which probably derived from three different lineages, found in Tahiti, Mo'orea and in the cluster Raiatea–Huahine. We outline the taxonomic consequences of this phylogeny. Our results support the hypothesis that evolutionary pattern, not distance between islands, shaped the long-distance colonization of oceanic islands by reed warblers.     

Polygyny and the breeding success of the great reed warblerAcrocephalus arundinaceus

Settlement date, mating status, and breeding success of individually marked great reed warblers, Acrocephalus arundinaceus, were studied during the 1980–84 breeding seasons in Kahokugata, Ishikawa Prefecture, Japan. Twenty-five per cent of the territorial males were polygynous, of which the majority were bigamous. The settling periods of both sexes were long, extending for 65 days in males and 49 days in females. About 80% of males and females settled in the first half of the settling period, and the settlement date of 28–54% males overlapped with that of females. Many of the late settlers were bachelors and the males which mated earlier tended to be polygynous. The timing of a male's settlement is important in acquiring mates. Fifty-five per cent of eggs laid were lost before fledging, mainly due to predation. The mean number of fledglings was 3.19 per primary female, 2.41 per secondary female, and 2.80 per monogamous female. Comparison of the number of fledglings of females which mated during the same period showed that the presence of another female in the same territory did not adversely affect the breeding success of either of the polygynous females. Polygynous males have the advantage of decreasing the risk of breeding failure under high predation pressure.