Acrocephalus Arundinaceus Research Articles

European CuckooCuculus canorusparasitism and host's rejection behaviour in a heavily parasitized Great Reed WarblerAcrocephalus arundinaceuspopulation

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

Predictors of natal dispersal in great reed warblers: results from small and large census areas

For most species, it is currently not known whether individual characteristics and population parameters affecting dispersal distances within the local population are also affecting dispersal of individuals emigrating longer distances. We evaluated this in the great reed warbler Acrocephalus arundinaceus by analysing data of natal dispersal distances within (1) the local hatching area in southern Central Sweden and (2) the species’ whole breeding range in Sweden. The different scales gave conflicting results. Site of birth was the main predictor of natal dispersal distance in the local census area, whereas in the larger census area the dispersal distance differed significantly between hatching years. Previous studies have shown that the recruitment rate of great reed warblers within the local study area differs between hatching years and also that it declines during the course of the breeding season. It was not known, however, whether these relationships reflected variation in survival or dispersal. The results from the present study suggest that the differences in local recruitment rate between hatching years is partly caused by annual variation in emigration rate, whereas the decline in local recruitment rate with season mainly reflects mortality of late‐hatched nestlings. Our study shows a twofold advantage of a large census area when studying dispersal. First, predictors of longer‐distance dispersal will be revealed and, second, the variation in local recruitment rate can be separated into the two components, survival and dispersal.

Some Aspects of Laying, Incubation and Hatching in the Great Reed-Warbler

During the breeding season of 1998, breeding ecology of the Great Reed-Warbler (Acrocephalus arundinaceus orientalis) was studied at Yangsoo-ri and Yongdam-ri of the Yangpyung-gun, Kyunggi province, Korea. Egg-weight (CV: 6.25) was more variable than either length or breadth, and breadth was the least variable of the measures. Significant variations in overall egg-weight occurred between clutches, and that more of the total variation in egg-weight and shape are due to inter-clutch variation as to intra-clutch variation when the data were pooled. The last egg tends to be larger than the remaining eggs in the clutch of the Great Reed-Warbler, suggesting the Great Reed-Warbler may adopt the brood-survival strategy. When method 3 was used, the most common incubation period is 12 days. In the Great Reed-Warbler, the length of the incubation period was related to clutch-size when method 1 (r=0.485, p<0.05) and method 2 (r=0.621, p<0.01) were employed, but not related to egg weight. The averagee number of days of hatching asynchrony was 2.5, raging 0.5~2.5. Asynchronous hatching was related to the clutch size (r=0.66, p<0.01). Hatching sequence was closely related to the laying sequence (r=0.93, p<0.001), suggesting Great Reed-Warblers incubate their eggs before clutch completion. The effect of egg weight on hatching asynchrony was found in Great Reed-Warblers (t-test, p<0.01).

Breeding ecology of the Eastern Great Reed Warbler Acrocephalus arundinaceus orientalis at Lake Kasumigaura, central Japan

CapsuleTwo populations of the Eastern Great Reed Warblers, one close to settlements and one far away, differed mainly in nest losses which were higher at the latter site. Aims To compare the breeding ecology of two Japanese populations, and with breeding data from Polish sites. Methods We studied two sites in 1996. The Seimei site comprises a natural reed-bed (Phragmites australis) (2.0 ha) c. 2 km from residential areas. The Sakura site occurs as smaller reed-bed (0.8 ha) close to residential areas. Field study comprised mainly nest searching and nest monitoring. Results Breeding density, nest height, number of stems supporting nest, clutch size and brood size were not significantly different between sites. The proportion of nests built on reed stems standing in the water (60%) was significantly higher at the Seimei site (P < 0.001), and the warblers in this habitat experienced significantly higher nesting failure (P < 0.05). Predation was a principal cause of failure at both sites. Significantly more nests suffered from predation at Seimei (53%) than at Sakura (16%). Breeding density was higher at both sites compared with European populations but it fell within ranges of other Japanese populations. Mean first-egg date of first brood does not differ between our sites and Polish ones. Conclusion The differences between the two Japanese populations and between Japanese and Polish populations were relatively small and were seen mainly in nest losses, breeding density and proportion of nests built over water.

Habitat selection of passerine birds nesting in the Ebro Delta reedbeds (NE Spain): Management implications

In this study, we explored the habitat associations of the five most abundant passerine species nesting in the Ebro Delta reedbeds (NE Spain). Habitat characteristics (water depth, vegetation height and density, and plant-species composition) and abundance of all passerine species were measured at each of the 68 sampling points. The sampled area included most of the reedbeds in the Ebro Delta. Sampling points were classified according to the habitat paramenters measured using cluster analysis. The resulting four habitat types differed in the composition and attributes of their passerine communities. The abundance of the great reed warbler (Acrocephalus arundinaceus) and reed warbler (Acrocephalus scirpaceus) was greatest in tall, permanently inundated and almost monospecific Phragmites reedbeds. In contrast, the numbers of the reed bunting (Emberiza schoeniclus) and Savi’s warbler (Locustella luscinioides) were greatest in areas of drier soils and a denser basal stratum. Finally, the fan-tailed warbler (Cisticola juncidis) showed no clear habitat association. In the Ebro Delta, reedbed management includes freshwater inputs and burning of standing reeds. These practices promote tall, permanently inundated reedbeds with minimal detrital accumulation. Bird species richness and diversity were greater in less intensely-managed habitats, but only diversity was significant (p<0.001). We conclude that, since reedbeds are important habitats for several scarce bird species, and the abundance of these species seem to be influenced by vegetation structure, bird conservation considerations should be incorporated into future reedbed management practices.

Restricted dispersal in a long-distance migrant bird with patchy distribution, the great reed warbler.

In patchily distributed species dispersal connects local populations into metapopulations. Reliable quantifications of dispersal are therefore crucial to understanding the population dynamics and genetic structure of such metapopulation systems. The great reed warbler (Acrocephalus arundinaceus) inhabits eutrophic lakes and has a patchy breeding distribution. In this study we investigated the dispersal pattern of the great reed warbler based on an extensive capture-recapture effort covering a large census area (22,500km2). At two adjacent breeding sites (10km apart) in southern Central Sweden, the "main study area", we ringed the majority of adult and nestling great reed warblers between 1992 and 1999. In 1998 and 1999, we opportunistically searched for territorial males at the majority of the Swedish breeding sites, and were able to examine about 56% of all males in the region. Analyses of recaptured males demonstrated that philopatry predominated. Sixty-nine percent of the recruiting nestlings returned to breed in the main study area (their natal area), and 92% of the resighted adults were found at the same breeding locality in both study years. Breeding dispersal was significantly more restricted than natal dispersal. Additional data on natal and breeding dispersal within the main study area in 1992-1999 suggested that females were as philopatric as males. The overall high level of philopatry, with only occasional longer dispersal distances documented, yielded a root-mean-square dispersal distance of 33km per generation. High philopatry, short dispersal distances and similar dispersal patterns of male and female great reed warblers contrast the findings among birds in general, but conform to data of species having patchy breeding habitat and isolated populations. Restricted dispersal suggests limited gene flow even among several Swedish populations, which is in line with some previous findings of the population ecology of the great reed warbler: (1) structured mtDNA lineages among European populations; (2) small-scale population differences in song patterns; and (3) low genetic variation and occurrence of inbreeding depression in our main study population.

Effect of winter cutting on the passerine breeding assemblage in French Mediterranean reedbeds

Common reed is increasingly harvested from the Mediterranean region toprovide thatching material to north European countries. The impact of thesemanagement practices on the fauna is poorly known. The aim of this study was toquantify the effect of reed cutting in the Mediterranean region through acomparative analysis of water regime, vegetation structure, arthropoddistribution and passerine assemblage at cut and uncut reedbeds in southernFrance. Cut reedbeds were characterised by a lower salinity, higher water levelin spring, and higher reed biomass than uncut reedbeds. Arthropod distributiondiffered consistently between cut and uncut sites, leading to a higher index offood available to passerines in cut reedbeds. Cut reedbeds had a similar birdspecies richness but a lower bird abundance, due to the significant decrease inMoustached Warblers and Bearded Tits at cut sites. The mild Mediterranean winterfavoured early growth of reed in spring, making harvested reedbeds suitable forbreeding of long-distance migrants such as the Great Reed Warbler and ReedWarbler. However, for the resident species that breed earlier in the season, cutreedbeds presumably lack sufficient vegetation cover to provide adequate nestingand feeding sites. Although biennial cutting (double wale) is considered as agood compromise between conservation and commercial interests in the UK, thejuxtaposition of annually cut and never cut reed patches appears as the onlysustainable alternative for the Mediterranean region. We further hypothesisethat an optimal mosaic design of cut/uncut reed patches could provide as high aconservation value as unmanaged reedbeds.

ADAPTATIONS BY GREAT REED WARBLERS TO BROOD PARASITISM: A COMPARISON OF POPULATIONS IN SYMPATRY AND ALLOPATRY WITH THE COMMON CUCKOO

Abstract Egg rejection behaviour, as an antiparasite defence against brood parasitism was compared in two great reed warbler Acrocephalus arundinaceus populations. In central Hungary the host population lives in sympatry with the common cuckoo Cuculus canorus, and it is heavily parasitised (ca 66%). In central Greece the great reed warbler occurs in allopatry with the cuckoo, so it is not parasitised at all. Experimental parasitism with poorly mimetic (often called 'non-mimetic') artificial cuckoo eggs revealed a 100% rejection rate towards the foreign eggs in the Greek population, but the Hungarian hosts accepted 29.4% and rejected 70.6% of the model cuckoo eggs. No other rejection type than ejection was observed. The significantly higher level of rejection in Greece than in Hungary suggests that the Greek great reed warbler population is an abandoned host population of the cuckoo, which kept its egg recognition ability. A computer-based image analysis of host's clutches revealed that interclutch variation in egg colour was greater than intraclutch variation in the Hungarian population, but not in the Greek population. Spottedness of eggs were similar in both populations. Generally, a tendency for a higher interclutch variation was found in Hungary. Intraclutch variation was more similar in the two populations, but for the colour variables it showed a lower value in Hungary than in Greece. The heavy cuckoo parasitism in Hungary is suggested to be the main cause for the increased interclutch variation in relation to intraclutch variation in egg appearance among great reed warbler clutches. The high interclutch variation reduces the chance that the parasite egg's appearance matches that of the hosts', which facilitates hosts' egg discrimination.

How to hatch from an egg of great structural strength. A study of the Common Cuckoo

Brood parasitism represents a unique mode of avian reproduction that requires a number of adaptations. For example, to reduce chances of puncture ejection of their eggs by small hosts, brood parasites may have been selected for laying eggs of unusually great structural strength. However, great structural strength of eggshells should hinder hatching. The goals of our study were to establish if chicks of the Common Cuckoo Cuculus canorus have more difficulty with hatching out of their strong eggs than chicks of species with eggs of similar size, and whether they possess any mechanisms facilitating hatching. To achieve these goals, we compared hatching pattern and selected body characteristics of chicks of the Common Cuckoo with those of another altricial species with eggs of a similar size, the Great Reed Warbler Acrocephalus arundinaceus. Although the rate of pecking was similar in the two species, the Common Cuckoo chicks started pecking earlier in relation to their emergence and consequently required more time and a greater cumulative number of pecks for breaking open their eggs than did young Great Reed Warblers. The two species also differed with respect to the pattern of opening their shells; in contrast to the warbler chicks, which enlarged the original pip circularly, the cuckoo chicks opened the egg by systematically creating a long narrow slit until they emerged. Finally, our study of hatched young revealed several differences; the Cuckoo hatchlings were significantly heavier, had a longer forearm, and their egg tooth was located significantly farther from the tip of the beak. The edge used for cutting through the shell was also significantly longer than that of hatchling Great Reed Warblers. To conclude, our data suggest that hatching is more difficult for a Cuckoo than for a Great Reed Warbler and that Cuckoos possess several mechanisms to overcome the problems of hatching from a structurally strong egg.

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.

Egg Mimicry in Cuckoos Parasitizing Four Sympatric Species of Acrocephalus Warblers

The Cuckoo (Cuculus canorus) is an interspecific brood parasite that mimics the eggs of its hosts. In this study we tested the resemblance between Cuckoo and host eggs in four host species breeding in sympatry: the Reed Warbler (Acrocephalus scirpaceus), Great Reed Warbler (A. arundinaceus), Sedge Warbler (A. schoenobaenus), and Marsh Warbler (A. palustris). According to the “gentes theory,” individual Cuckoos lay eggs that mimic those of a single host species, and they parasitize only that species. We therefore expected the Cuckoo eggs to more closely resemble the eggs of their respective host species than eggs of sympatric hosts. However, analyses showed no such resemblance; test subjects were not able to distinguish Cuckoo eggs taken from nests of different host species. The most common Cuckoo egg type resembled the eggs of the Garden Warbler (Sylvia borin), a frequent European Cuckoo host which, however, has not been found parasitized in the study area. A possible evolutionary origin for this resemblance is discussed. Results from radio-tracking of Cuckoo females in the study area have shown that host preference does exist. This preference has not yet been manifested in the evolution of clear Cuckoo egg morphs adapted to the different host species. Imitación de Huevos por Cuculus canorus que Parasitan Cuatro Especies Simpátricas de Acrocephalus Resumen. El cuco (Cuculus canorus) parasita los nidos de varias especies imitando los huevos de sus especies hospederas. En este estudio cotejamos la similitud entre los huevos de cucos y los huevos de cuatro especies hospederas que crían en simpatría: Acrocephalus scirpaceus, A. arundinaceus, A. schoenobaenus, A. palustris. De acuerdo a la “teoría de gentes” los huevos que pone un individuo de cuco imitan sólo a los huevos de una especie hospedera, y por lo tanto parasita sólo a esa especie. Debido a esto esperábamos que los huevos de cucos fueran más similares a los huevos de sus respectivas especies hospederas que a los huevos de las demás especies hospederas simpátricas. Sin embargo, los análisis no mostraron dicha similitud; los individuos testeados no fueron capaces de distinguir entre huevos de cucos tomados de nidos de diferentes especies hospederas. El tipo de huevo más común de cuco se asemeja a los huevos de Sylvia borin, una especie hospedera frecuente para los cucos europeos, la cual, sin embargo, no ha sido encontrada parasitada en el área de estudio. Se discute el posible origen evolutivo de dicha similitud. Los resultados de telemetría de hembras de cucos en el área de estudio han mostrado que la preferencia de hospedadores existe. Esta preferencia no se ha manifestado aún en la evolución del morfo claro de huevo de cuco adaptado a las diferentes especies hospederas.

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.

Increase of genetic variation over time in a recently founded population of great reed warblers (Acrocephalus arundinaceus) revealed by microsatellites and DNA fingerprinting.

Genetic similarity within pairs of individuals was examined using both 10 polymorphic microsatellite loci and multi-locus DNA fingerprinting profiles in a semi-isolated population of great reed warblers at Lake Kvismaren, south Central Sweden, in 1987-1993. The population was founded by a few individuals in 1978, followed by a gradual increase in numbers until 1988, since when the population has remained relatively stable with about 60 breeding birds. We have previously found that high genetic similarity between pair-mates in the population during the early part of the study period reduced egg hatching success, and hence reproductive success. The measures of pairwise genetic similarity, microsatellite allele sharing and DNA fingerprinting band sharing, were highly correlated with pedigree-based relatedness. Both microsatellite and DNA fingerprinting similarities between pair-mates declined significantly over the study period, and the pattern was most pronounced in the DNA fingerprinting data. Analyses restricted to the microsatellite data showed that the average annual microsatellite similarity between pairwise combinations of individuals, as well as individual homozygosity in males, declined significantly over the study period, and that several immigrants carrying novel alleles entered the population during the study. Hence, the temporal decline in genetic similarity of mates in the population is probably a consequence of increased immigration, facilitated by the recent expansion of the species in the region. These results suggest that the population has now recovered genetically, or is in the process of recovering, from a recent founder event.

The quality and the timing hypotheses evaluated using data on great reed warblers

The seasonal decline in reproductive success observed in many animal species may be caused by timing per se (timing hypothesis) or by variation in phenotypic quality between early and late breeding females (quality hypothesis). To distinguish between these two hypotheses, several studies of birds have used clutch removal experiments to manipulate breeding date. However, removal experiments also increase the females’ previous reproductive effort due to the production of an extra clutch and a longer incubation period. According to life‐history theory an increase in reproductive effort lowers future reproduction. Hence, life‐history theory predicts lowered success of replacement broods for other reasons than expected from the timing hypothesis. Female great reed warblers, Acrocephalus arundinaceus, studied in Sweden are frequently exposed to nest predation, after which many lay replacement clutches. In order to examine possible effects of previous reproductive effort on different fitness components, we analysed the re‐laying frequency and the reproductive success of replacement broods in relation to time of the season and previous reproductive effort (measured as the length of the previous breeding attempt, LPB). In clutches of re‐laying females both the number of fledglings and the proportion of recruits were negatively correlated with LPB, whereas re‐laying frequency and clutch size were not related to LPB. We expect such relationships to be present also among other species. Consequently, the use of replacement clutches, as for example in clutch removal experiments, in evaluations of the cause of the often observed seasonal decline in various fitness components, might exaggerate the importance of the timing hypothesis over the quality hypothesis.

Potential food resources and nestling food in the Great Reed Warbler (Acrocephalus arundinaceus arundinaceus) and Eastern Great Reed Warbler (Acrocephalus arundinaceus orientalis)

The main ecological difference between the Great Reed Warbler (Acrocephalus arundinaceus arundinaceus) and Eastern Great Reed Warbler (Acrocephalus acrocephalus orientalis) lies in the density of breeding population, which in the Eastern subspecies studied in Japan is on average 10 times higher than that of European populations of the Western subspecies. In this study it is shown that potential food resources of the Great Reed Warbler in Japan are more than ten times more abundant than those in Poland, with respect both to the number of potential prey items (invertebrates) and to their total biomass. This strongly suggests that such high densities of Great Reed Warbler in Japan are at least partly due to more plentiful food supply. Other factors which could explain the difference between breeding densities in Japan and Poland are the predation by Marsh Harrier (Circus aeruginosus) in Poland and lack of competition with Reed Warbler (Acrocephalus scirpaceus) in Japan. The abundance of food on the sample plot in Japan resulted primarily from outbreaks of dipterans of the suborder Nematocera, the most common of which were Chironomidae. Although Chironomidae were not the preferred prey, they made up a substantial proportion of the diet of nestlings. In both studied areas, apart from dipterans, the greatest proportion of nestlings' diet constituted arachnids. Differences in nestlings' diet between study sites resulted mainly from a greater proportion of Nematocera and lower proportion of Coleoptera and Odonata in Japan.

Effect of nest and nest site characteristics on the risk of cuckooCuculus canorusparasitism in the great reed warblerAcrocephalus arundinaceus

The risk of cuckooCuculus canorusparasitism for great reed warblerAcrocephalus arundinaceusnests was evaluated in respect to nest and nest site structure in the reedbeds along canals in central Hungary, In total 90 nests were analysed, from which 31 remained unparasitised. 37 were single parasitised and 22 multiple parasitised. Multiple discriminant analysis, based on eight structural variables, separated well the unparasitised clutches from the cases of single and multiple parasitism, but the latter two categories also showed a weaker separation. Nests close to cuckoo vantage points are most vulnerable to parasitism. The variable “distance from nest to closest available cuckoo perch site” (tree or electric wire) played the most important role in the risk of parasitism. It showed highly significant differences among the groups: lowest distances were found in the case of multiple parasitism, and the highest distances were measured when nests remained unparasitised. Additionally, “nest visibility” also showed significant differences among the three groups: a more visible nest is more likely to be parasitised. Risk of parasitism affects on the host on two levels: 1) female cuckoos search for nest‐building hosts from a perch site, but 2) when they are in the act of parasitism, they can find more visible nests in the reed. Besides the robust effects of the variables “distance to cuckoo perch site” and “nest visibility”, the variable “distance from nest to open water” may reduce, but “nest volume” increases the risk of multiple parasitism. Differences between cases of single and multiple parasitism are weak, mainly affected by chance. We explain it by the high abundance of the cuckoo, the parasitism rate was 65.6% (59/90), Cuckoos parasitised almost all of the available nests in the close vicinity of potential perch sites. There was no evidence that great reed warblers nested closer to each other when risk of parasitism was high.

Effect of nest and nest site characteristics on the risk of cuckoo Cuculus canorus parasitism in the great reed warbler Acrocephalus arundinaceus

Effect of nest and nest site characteristics on the risk of cuckoo Cuculus canorus parasitism in the great reed warbler Acrocephalus arundinaceus

Brood sex ratios, female harem status and resources for nestling provisioning in the great reed warbler ( Acrocephalus arundinaceus )

The theory of parental investment and brood sex ratio manipulation predicts that parents should invest in the more costly sex during conditions when resources are abundant. In the polygynous great reed warbler, Acrocephalus arundinaceus, females of primary harem status have more resources for nestling provisioning than secondary females, because polygynous males predominantly assist the primary female whereas the secondary female has to feed her young alone. Sons weigh significantly more than daughters, and are hence likely to be the more costly sex. In the present study, we measured the brood sex ratio when the chicks were 9 days old, i.e. the fledging sex ratio. As expected from theory, we found that female great reed warblers of primary status had a higher proportion of sons in their broods than females of lower (secondary) harem status. This pattern is in accordance with the results from two other species of marsh-nesting polygynous birds, the oriental reed warbler, Acrocephalus orientalis, and the yellow-headed blackbird Xanthocephalus xanthocephalus. As in the oriental reed warbler, we found that great reed warbler males increased their share of parental care as the proportion of sons in the brood increased. We did not find any difference in fitness of sons and daughters raised in primary and secondary nests. The occurrence of adaptive sex ratio manipulations in birds has been questioned, and it is therefore important that three studies of polygynous bird species, including our own, have demonstrated the same pattern of a male-biased offspring sex ratio in primary compared with secondary nests.

Nestling provisioning in polygynous great reed warblers ( Acrocephalus arundinaceus ): do males bring larger prey to compensate for fewer nest visits?

Most studies of social polygyny in birds have examined male provisioning on the basis of the number of feeding visits. This may be misleading if males compensate for infrequent visits by bringing larger prey at each visit. We investigated nestling provisioning in the socially polygynous great reed warbler, Acrocephalus arundinaceus, in south Central Sweden in 1996–1997. We collected data on rate of feeding visits, prey size and the amount of biomass delivered by males and females. Males had lower rates of feeding visits and provided smaller prey to nestlings in secondary than in monogamous and primary nests. Secondary females had higher rates of feeding visits and brought larger prey than monogamous and primary females. These results confirm that secondary females face a potential cost of polygyny through a lower rate of male feeding, and that this cost was reinforced by the significantly lower male provisioning rate (biomass h–1) at secondary nests. Secondary females compensated for the lack of male assistance by increasing their rate of feeding and bringing larger prey. As a result, offspring in nests of secondary females received as much food as did those in nests of primary females. Prey load size increased with the parent’s proportion of feeding visits, suggesting that parents use different feeding strategies depending on their amount of responsibility for nestling provisioning. We suggest that parents which take the main responsibility for nestling feeding have to forage further away from the nest, and based on optimal-foraging theory, they should then on average bring larger prey to their nest.

Patterns of Nest Predation Contribute to Polygyny in the Great Reed Warbler

According to the polygyny threshold model, females are compensated for the cost of sharing a territory with other females by breeding in territories of higher quality than those of monogamously mated females. In the polygynous Great Reed Warbler (Acrocephalus arundinaceus), the variation in territory quality may be associated with nest site characteristics or food supply. In this study, we examined the probability of nest predation in Great Reed Warblers in relation to an indirect measure of territory quality (territory attractiveness rank as indexed by settlement order) and to several variables of nest site structure. Artificial nests with attached soft plasticine model eggs were placed in Great Reed Warbler territories of different attractiveness. Marks in the soft model eggs were compared to marks from possible predator species. Both inter- and intraspecific predators were found to prey upon eggs in artificial nests. Marsh Harriers (Circus aeruginosus) predominantly preyed upon artificial nests located in territories of low attractiveness, in low-density reed beds, and early in the season. Bitterns (Botaurus stellaris) and Water Rails (Rallus aquaticus) showed a preference for artificial nests located inside the reeds. When all types of predators of artificial nests were pooled, the probability of nest predation was lower in more attractive territories. This pattern was corroborated in an analysis of predation on real nests. In general, polygyny occurred in territories with a low rate of nest predation, whereas unmated males held territories where the risk of predation was high. The low rate of nest predation in attractive territories on average compensated secondary females for the cost associated with sharing a male (i.e., reduced male assistance when feeding nestlings). The polygyny threshold model has been around for several decades, however, only few studies have identified the factors that may compensate secondary females for the cost of sharing a mate. In the present study we were able to show that the patterns of nest predation can be an important territory quality parameter that compensate secondary females, and, hence, contribute to the occurrence of avian polygyny.