Seasonal population changes of five parasitoids attacking the scale insect Nipponaclerda biwakoensis on the common reed, with special reference to predation by wintering birds

Seasonal changes in the abundance of five species of hymenopterous parasitoids (four species of Encyrtidae and one species of Eulophidae) attacking the scale insect Nipponaclerda biwakoensis on the common reed were investigated for 2 years in Lake Biwa, with special reference to predation by the reed bunting, Emberiza schoeniclus, during winter. The scales settled on reed shoot stems under sheath leaves, passing through three discrete generations per year. The abundance of adult female scales increased exponentially from July (first generation) to December (third generation). Adult female scales of the third generation overwintered on reed shoots. During winter, female scale abundance dramatically declined, whereas the number of predation marks made by reed buntings using their bills on reed sheath leaves increased. The generations of all five parasitoids were synchronized with the host scale generations, and the five parasitoids overwintered as larvae inside the scale bodies. The abundance of parasitized scales and parasitoid adults emerging from the scales also increased from July to December, but greatly decreased during winter. The overall parasitism rate of the female scales remained at relatively low levels (less than 40%) throughout the year, including before and after winter. A bird exclusion experiment revealed that the dramatic winter decrease of the abundance of the scale and its five parasitoids was due to intensive and non‐selective predation by the buntings on unparasitized and parasitized scales. Additionally, the proportion of immature parasitoids removed by birds varied between the five parasitoid species. Thus, seasonal population changes of the five scale parasitoids are considerably affected by bird predation on overwintering immature parasitoids.     

How female reed buntings benefit from extra-pair mating behaviour: testing hypotheses through patterns of paternity in sequential broods

Extra-pair paternity is an important aspect of reproductive strategies in many species of birds. Given that in most species females control whether fertilization occurs, they are expected to benefit in some way from the extra-pair matings. In this study we use patterns of extra-pair paternity (EPP) in broods of individual reed buntings (Emberiza schoeniclus), both within and between seasons, to test four hypothesized female benefits: (1) assessing potential future partners and seeking (2) genetic diversity (3) good genes, or (4) compatible genes. Reed buntings are socially monogamous, multibrooded passerines with extremely high levels of extra-pair paternity. We studied a population of reed buntings in the Netherlands in 2002 and 2003; 51% of offspring in 74% of nests were extra-pair. We showed that patterns of EPP did not support the first and second hypotheses, since females did not form a pair with previous extra-pair partners, EPP was not evenly distributed among broods and more broods than expected were sired by a single male. Furthermore, there was no relation between a male's within- and extra-pair fertilization success, no consistency in EPP between breeding attempts, no effect of parental relatedness on EPP and several cases of reciprocal paternity. These patterns do not support the good genes hypothesis and are most consistent with the genetic compatibility hypothesis. However, our previous finding that older males are more successful in gaining EPP, suggests some effect of good genes. These hypotheses need not be mutually exclusive, as females may select compatible males above a certain quality threshold (e.g. old males).     

Substantial contribution of invertebrates to the diet of a winter seed‐eater,the reed bunting Emberiza schoeniclus,wintering in a sewage farm in south‐western Poland

We investigated whether the winter diet of a typical seed‐eating bird is hard‐wired in the context of evolutionary hypothesis for granivory. We examined the diet composition of ‘a small‐billed form’ of the reed bunting Emberiza schoeniclus wintering in a sewage farm in south‐western Poland (Central Europe), where unfrozen wastewater provides various groups of invertebrate prey. The analysis of droppings (N = 151) collected from four different feeding grounds located in reedbeds and grasslands inundated with waste‐water showed the substantial contribution of invertebrates in the diet of reed buntings. Across four sample areas, the frequency of invertebrates in faecal samples ranged between 37% to 80%. In total, we identified 194 animal prey, mainly spiders Araneae (53% of all identified invertebrate prey), and several taxa of Coleoptera (43%). Among plant food (N = 8357 identified items), the most numerous were shells of weed seeds, namely Amaranthus sp. (56.8%), Urtica dioica (22.6%), and Chenopodium sp. (19.6%). Our results showed that, during winter, the reed bunting is not an obligatory seed‐eater. This species may exploit both animal and plant food; hence, our results indicate that the reed buntings wintering in temperate Europe may feed more opportunistically than was previously assumed. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 429–433.     

Ten microsatellite loci for evolutionary research on Nesospiza buntings

We tested 19 loci originally developed for Vidua and Geospiza, of which six Vidua loci were polymorphic and furthermore, developed four microsatellite loci for Nesospiza buntings. Allelic variation ranged from two to 13 alleles, with heterozygosity from 0 to 0.82. Ascertainment bias was evident as increased allelic diversity of loci developed for Nesospiza, versus those developed for Vidua. Two loci showed linkage, and several, deviation from Hardy–Weinberg equilibrium. We suggest such deviations are through introduction of new alleles by introgression, or of the Wahlund effect. These loci will contribute to the understanding of speciation in Nesospiza buntings of the Tristan da Cunha islands.     

Characterization of 38 microsatellite loci in the European blackbird, Turdus merula (Turdidae, AVES)

We characterized 38 microsatellite loci in the European blackbird, Turdus merula. Thirty-seven loci were identified by testing 242 loci that had been originally isolated in other avian species. One additional locus was isolated from a European blackbird genomic library. All loci were characterized in 20-29 blackbirds from a population in the Czech Republic and displayed between two and 16 alleles, with observed heterozygosity ranging from 0.04 to 1.00. Thirty-seven loci could be assigned a chromosome location in the zebra finch (Taeniopygia guttata) genome based on sequence homology.     

Diversification at high latitudes: speciation of buntings in the genus Plectrophenax inferred from mitochondrial and nuclear markers

High‐latitude diversification is a process characterized by speciation and extinction due to climatically driven vicariance and dispersal events. McKay’s buntings (Plectrophenax hyperboreus) are high‐latitude island endemic songbirds, and their global range is restricted to Beringia. Snow buntings (P. nivalis), their closest relatives, are distributed throughout the Holarctic, breeding in available habitat surrounding the island range of McKay’s buntings. We sequenced 1123 base pairs of mitochondrial DNA for 40 individuals of each species and analysed a total of 913 AFLPs for 57 individuals. Both marker types suggested weak but significant genetic differentiation. Analysis of sequence data indicated divergence occurring when the current breeding range of McKay’s buntings was a hill on the Beringian steppe (～18 400 to ～73 700 years before present), suggesting that snow buntings were restricted to lower latitudes by ice sheets. Ancestral effective population size estimates indicate a founder event in McKay’s buntings followed by an expansion and then a reduction in effective size. Rising sea levels and asymmetric hybridization from McKay’s buntings into the postglacially‐colonizing population of snow buntings could account for this reduction. Reproductive isolation is likely maintained through differential arrival dates on breeding grounds and the high breeding density of McKay’s buntings. This recent, high‐latitude divergence best fits a model of founder event speciation driven by vicariance and oscillations in habitat due to climate change.     

Characterization of 20 microsatellite loci in the long‐tailed tit Aegithalos caudatus (Aegithalidae,AVES)

We characterized 20 microsatellite loci in the long‐tailed tit, Aegithalos caudatus. Polymorphic loci were identified by testing 114 loci that had been originally isolated in other avian species. The loci were characterized in 23–163 unrelated long‐tailed tits from a British population and displayed between two and 42 alleles, with observed heterozygosity ranging between 0.09 and 0.97.     

Isolation and characterization of microsatellite markers in Dendrobium officinale,an endangered herb endemic to China

As an invaluable herb, Dendrobium officinale has been in severe danger since 1950 because of human exploitation and habitat deterioration. In order to investigate the genetic diversity and structure of this species, we isolated and characterized 12 microsatellite loci derived from two microsatellite‐enriched libraries. Twenty‐two individuals from Leye population were analysed. These loci were polymorphic and displayed three to 12 alleles per locus. The observed and expected heterozygosities ranged from 0.150 to 0.624 and from 0.162 to 0.605, respectively. These are the first microsatellite loci characterized from D. officinale that will contribute to research on the conservation, genetic diversity, population structure and individual authentication.     

Part-Time Mate Guarding Affects Paternity in Male Reed Buntings (Emberiza schoeniclus)

Male mate guarding by close following has been reported in many socially monogamous bird species and is generally believed to function as a paternity guard. Many aspects of the dynamics and effectiveness of this behavior are still however poorly understood. Here, we describe the temporal variation in mate guarding behavior in male reed buntings (Emberiza schoeniclus) with a particular focus on how males allocate their mating effort between mate guarding and extrapair mating in a context of intense sperm competition. In our highly synchronous study population most males have to balance the simultaneous and mutually exclusive demands of mate guarding and seeking extrapair copulations (EPCs). We found that males frequently switched between guarding their mates and performing intrusions to neighboring territories. Both activities seemed to have significant fitness payoffs, as male mate guarding effort had a positive effect on paternity, and a large fraction of extrapair fertilizations occurred during the days when the sire guarded its own female. The reed bunting is thus an example of how discontinuous or part‐time mate guarding can still be effective in securing paternity. Female reed buntings were not particularly active in initiating EPCs as they never were observed performing extraterritorial forays. We argue that the absence of female‐initiated EPCs is a prerequisite for males to trade mate guarding against seeking EPCs. Otherwise, if females circumvent male mate guarding by timing their EPCs to periods of male absence, males should guard their mates almost continuously or rely on alternative paternity guards.     

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.     

Characterization of Mauritius parakeet (Psittacula eques) microsatellite loci and their cross-utility in other parrots (Psittacidae, Aves)

We characterized 21 polymorphic microsatellite loci in the endangered Mauritius parakeet (Psittacula eques). Loci were isolated from a Mauritius parakeet genomic library that had been enriched separately for eight different repeat motifs. Loci were characterized in up to 43 putatively unrelated Mauritius parakeets from a single population inhabiting the Black River Gorges National Park, Mauritius. Each locus displayed between three and nine alleles, with the observed heterozygosity ranging between 0.39 and 0.96. All loci were tested in 10 other parrot species. Despite testing few individuals, between seven and 21 loci were polymorphic in each of seven species tested.     

Microsatellite DNA markers for population-genetic studies of Labeo dyocheilus (McClelland, 1839)

Fifty‐four primers published for six cyprinid fishes were tested to amplify homologous microsatellite loci in Labeo dyocheilus. Fifteen primers yielded successful amplification and seven were polymorphic with 3–9 alleles. To evaluate utility of the identified loci in population genetic study, 84 samples were analysed. The samples were collected from four rivers viz. Beas, Satluj, Yamuna and Jiabharali. The four sample sets displayed different diversity levels, with observed heterozygosity from 0.34 to 0.53. Significant genotype heterogeneity (P < 0.001) over all loci indicated that the samples are not drawn from the same genepool. The identified microsatellite loci are promising for use in fine‐scale population structure analysis of L. dyocheilus.     

Fragmentation within and between wetland reserves: the importance of spatial scales for nest predation in reed buntings

Wetlands in many parts of the world are affected by fragmentation at multiple spatial scales. In Switzerland, most wetlands have been destroyed over the past two centuries and management of the remaining wetland reserves has intensified in the recent years leading to increased fragmentation of reed areas within reserves. Using four years of data on the reproductive performance of color-banded reed bunting Emberiza schoeniclus populations, we explored fragmentation effects on nest predation rates at four spatial scales ranging from the nest to the landscape scale. In the egg stage, predation rate was negatively related to vegetation cover, vegetation height and nest height, but positively linked to water cover and depth next to the nest (nest scale). Probability of predation declined with increasing size of reed patches containing the nests and distances of nests to the water and land sided reed edge, as well as with decreasing edge to area ratio (edge scale). There was a weak positive association between degree of fragmentation of reed patches within sites and nest predation rates (site scale). Finally, nest predation probability increased with distance to the nearest wetland (landscape scale). Jointly analyzing variables from different spatial scales revealed that a model combining variables from the nest, edge and landscape scale best explained predation probability in the egg stage. In the nestling stage, the single most important factor influencing nest predation probability was the distance to the nearest wetland (landscape scale), with nest predation decreasing with distance between sites. Our results show that the probability of nest predation in reed buntings is affected by fragmentation within and between wetland reserves and that the effects differ between breeding stages. Future management of wetland reserves should aim at sparing reed patches large and dense enough to provide safe nest sites for birds.     

Small and large wetland fragments are equally suited breeding sites for a ground-nesting passerine

Large habitat fragments are generally thought to host more species and to offer more diverse and/or better quality habitats than small fragments. However, the importance of small fragments for population dynamics in general and for reproductive performance in particular is highly controversial. Using an information-theoretic approach, we examined reproductive performance and probability of local recruitment of color-banded reed buntings Emberiza schoeniclus in relation to the size of 18 wetland fragments in northeastern Switzerland over 4 years. We also investigated if reproductive performance and recruitment probability were density-dependent. None of the four measures of reproductive performance (laying date, nest failure probability, fledgling production per territory, fledgling condition) nor recruitment probability were found to be related to wetland fragment size. In terms of fledgling production, however, fragment size interacted with year, indicating that small fragments were better reproductive grounds in some years than large fragments. Reproductive performance and recruitment probability were not density-dependent. Our results suggest that small fragments are equally suited as breeding grounds for the reed bunting as large fragments and should therefore be managed to provide a habitat for this and other specialists occurring in the same habitat. Moreover, large fragments may represent sinks in specific years because a substantial percentage of all breeding pairs in our study area breed in large fragments, and reproductive failure in these fragments due to the regularly occurring floods may have a much stronger impact on regional population dynamics than comparable events in small fragments. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Characterization of microsatellite loci In The hihi Notiomystis cincta (Notiomystidae, Aves)

We have characterized 20 polymorphic microsatellite loci in the hihi Notiomystis cincta. Loci were identified by testing loci originally isolated in other avian species and by isolating new microsatellites from a hihi genomic library. These loci were characterized in unrelated hihi from a single population on Tiritiri Matangi Island (n = 98). Each locus displayed between two and 10 alleles, and observed heterozygosities ranged between 0.29 and 0.91. Nineteen of the 20 polymorphic loci could be assigned a chromosome location in the zebra finch, Taeniopygia guttata genome based on sequence homology.     

Isolation and characterization of microsatellite markers from reed parrotbill, Paradoxornis heudei (Aves: Timaliidae)

Eleven microsatellite markers were obtained from reed parrotbill, Paradoxornis heudei, using the fast isolation by AFLP of sequences containing repeats (FIASCO) method as part of an effort to compare levels of genetic diversity in different populations of China. Polymorphism levels ranged form 5 to 11 alleles (mean = 9.27) using 32 reed parrotbills, with the observed heterozygosity ranging from 0.037 to 0.80. Six loci were significant deviated from HWE and nine loci showed linkage equilibrium. The utility of these loci on two other Passeriformes species, vinous-throated parrotbill (P. webbianus) and oriental great reed warbler (Acrocephalus orientalis), were also tested.     

PERMANENT GENETIC RESOURCES: Isolation and characterization of polymorphic trinucleotide and dinucleotide microsatellite loci for the ice goby, Leucopsarison petersii

Eleven microsatellite markers were isolated and characterized for the ice goby, Leucopsarion petersii, from genomic libraries enriched for (ATG)(9) and (CA)(16) . Twenty individuals from a single population were used to screen polymorphism in these loci. The number of alleles per locus and observed heterozygosity ranged from four to 22 and from 0.35 to 1.00, respectively. All loci did not significantly deviated from Hardy-Weinberg equilibrium, and there was no evidence of linkage disequilibrium between all loci-pairs. These loci showed Mendelian inheritance in a full-sib family. The high level of polymorphism of these loci will be useful for studies of population genetics.     

Isolation, characterization and chromosome locations of polymorphic black-billed magpie Pica pica (Corvidae, AVES) microsatellite loci

One-hundred and two unique loci were isolated from a microsatellite-enriched black-billed magpie Pica pica genomic library. Sixteen of these new loci, along with 148 existing microsatellite passerine loci, were tested for polymorphism in four unrelated black-billed magpies. We identified a total of 43 unique polymorphic loci (10 and 33 respectively) that we characterized in 24 unrelated individuals from a population in Guadix, Spain. The putative chromosomal locations of loci polymorphic in black-billed magpie were assigned based on the location of their orthologues in the assembled zebra finch and chicken genomes.     

Isolation and characterization of microsatellite loci in the western long-fingered bat, Miniopterus magnater

We isolated and characterized 10 microsatellite loci in the western long-fingered bat, Miniopterus magnater. These loci were tested on 48 individuals from Anhui Province of China, and all loci were highly polymorphic. The mean number of observed alleles per locus was 13.6 (range from six to 27). Observed and expected heterozygosity values ranged from 0.364 to 0.957, and from 0.676 to 0.951, respectively. After Bonferroni correction, four loci deviated significantly from Hardy-Weinberg equilibrium. No pairs of loci were in linkage disequilibrium. These polymorphic markers will be used to examine population structure and genetic diversity in this species.     

No evidence for increased offspring heterozygosity from extrapair mating in the reed bunting (Emberiza schoeniclus)

It has been hypothesized that females mate multiply to increasethe heterozygosity of their progeny because heterozygous individualsare assumed to have a fitness advantage. Females can maximizeheterozygosity of their offspring by mating with geneticallyunrelated and/or heterozygous males. We tested these predictionsin a socially monogamous passerine, the reed bunting (Emberizaschoeniclus), where extrapair paternity occurs frequently. Theresults based on genotypes at nine microsatellite loci revealedthat females were no less genetically related to the extrapairmale(s) (EPMs) than to their pair male (i.e., breeding partner)and that EPMs were no more heterozygous than the males theycuckolded. In addition, a direct comparison of maternal half-siblingsnaturally raised in the same brood showed that extrapair youngwere no more heterozygous than within-pair young. Thus, femalereed buntings do not seem to mate with EPMs to increase offspringheterozygosity. It is not yet known whether extrapair matinginvolves any benefits at all to females in this species.