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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Influences of wind flow on stopover decisions: the case of the reed warbler Acrocephalus scirpaceus in the Western Mediterranean

Wind directions measured at two different heights (850 hPa and 700 hPa) and at different hours of the night were analysed during the spring migration passage at a bird stopover site located in the western Mediterranean, in order to evaluate the importance of wind components for a stopover decision. From a huge ringing campaign of bird migration in north-east Spain, data from the reed warbler Acrocephalus scirpaceus have been used for the analysis. From a total of 2,478 reed warblers captured between 1993 and 1997 data recording significant days, with a high number of captures, and decrease days, with few captures, have been selected to develop an analysis of wind direction in relation to stopover and flight resumption. On days with a high capture the winds had mainly a fourth-quadrant flow (from the north, north-west and west), these being mainly head winds. Winds with westerly component (from the north-west, west and south-west), which enhance the flight, account for the majority of the days when there was a low capture of reed warblers. Wind direction therefore appears to be a determining factor for stopover decisions and resumption of flight for the reed warblers at an intermediate stage of their spring migration where topographical characteristics govern the winds.     

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.     

An exceptionally high diversity of hoverflies (Syrphidae) in the food of the reed warbler (Acrocephalus scirpaceus)

Despite being considered a classical example of protective Batesian mimicry hoverflies (Syrphidae) are known to be preyed upon by various passerines. The aim of the present study was to examine in detail food brought by reed warblers Acrocephalus scirpaceus to their nests to better understand the importance of hoverflies in the diet of small passerines. Using neck collars, 273 food samples containing 8,545 food items delivered to reed warbler and parasitic common cuckoo Cuculus canorus nestlings in warbler nests were recorded. The study was conducted during three breeding seasons in South Moravia, Czech Republic. An unusually high diversity of hoverflies was found — 27 species, including Mesembrius peregrinus (critically endangered species in the Czech Republic) and Mallota cimbiciformis (endangered species) — a new taxon to the Czech Republic. This indicates that nestling diet analyses may provide not only information on avian foraging behaviour but also important faunistic data. Thus, without the detailed identification to species level of material from foraging behaviour studies valuable scientific information may be lost. Overall dominance of Syrphidae was 3.7%, the most common species being Episyrphus balteatus (55.7%, n = 318). However, this number seriously underestimates the importance of hoverflies in the diet of reed warblers as hoverflies are one of the largest prey taken by warblers. Both larvae and pupae were rare, imagines strongly dominating (92.7%). Both specific wasp mimics (e.g., Chrysotoxum verrali) and bee mimics (e.g., Eristalis spp.) were not avoided by foraging reed warblers. The presence of a parasitic cuckoo chick did not affect host foraging behaviour with respect to overall dominance of hoverflies in the diet (warbler 3.3%, cuckoo 3.8%).     

Discovery of a breeding area of the enigmatic large‐billed reed warbler Acrocephalus orinus

The present study reports the discovery of the hitherto unknown breeding grounds of the large‐billed reed warbler Acrocephalus orinus in the Badakhshan region of Tajikistan. A total of eight adult individuals were caught at three riverine woodland sites and breeding was confirmed at one locality. The identity of the birds was confirmed on the basis of partial sequences of the cytochrome b gene. Even among individuals sampled at the same locality, genetic diversity was surprisingly high. This was interpreted as the result of secondary contact between previously isolated or fragmented populations. The breeding range of the large‐billed reed warbler is likely small and the species’ habitat is under intense pressure from the collection of firewood and fodder. Consequently, the large‐billed reed warbler remains of considerable conservation concern.     

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.     

Prey abundance and male parental behaviour in Acrocephalus warblers

This study examined the prey abundance at nesting sites of five different Acrocephalus warbler species. Intraspecific variations in prey density were analysed to determine the relative significance of this ecological factor for different mating systems and components of male parental behaviour. The Great Reed Warbler Acrocephalus arundinaceus, a facultatively polygynous species, nested in areas with the highest insect abundance. Its intraspecific variation in prey density (between different territories) was the highest while the male: female offspring feeding ratio was the lowest. At the other extreme, the monogamous Moustached Warbler Acrocephalus melanopogon and Reed Warbler Acrocephalus scirpaceus nested in areas with the lowest prey abundance and lowest intraspecific variation. Males of these species had the highest involvement in feeding young. In the Moustached Warbler and Marsh Warbler Acrocephalus palustris, an intraspecific comparison revealed a negative correlation between insect abundance and the male feeding component; a tendency in this direction was found for the Reed Warbler, while no correlation was evident in the polygynous Great Reed Warbler. These data demonstrate a general pattern relating prey abundance, mating systems and male parental investment on an interspecific level. However, intraspecific correlations revealed a net of complex interdependence. One of the factors which may be important is resource predictability. We found a significant correlation between certain vegetation types and insect abundance. The vegetation composition of territories is further influenced by interspecific dominance relationships whereby the dominant Great Reed Warbler occupies territories with the vegetation type which correlates with high prey abundance while the overlapping Moustached Warbler and Reed Warbler are restricted to areas with lower insect density.     

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.     

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.     

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.     

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.     

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.