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

Heritability of dispersal in the great reed warbler

Dispersal is commonly considered to be a condition-dependent behaviour with no or low heritability. Here, we show that dispersal in the great reed warbler (Acrocephalus arundinaceus ) has a high heritability. Analyses of capture-recapture data of male great reed warblers gathered from the species' whole Swedish breeding range revealed a remarkable offspring-parent resemblance in dispersal behaviour (philopatry vs. inter-population dispersal). Also, the degree of dispersal differed between cohorts, which shows that dispersal was partly conditionally dependent. The offspring to mid-parent estimate of heritability was 0.50. In a previous study of the same data set of male offspring, we did not detect associations between dispersal and several relevant environmental, parental and offspring condition factors. Thus, our results indicate that variation in dispersal partly has a genetic basis in great reed warblers

A new approach to study dispersal: immigration of novel alleles reveals female-biased dispersal in great reed warblers

We use the assignment technique and a new approach, the 'novel allele technique', to detect sex-biased dispersal in great reed warblers Acrocephalus arundinaceus . The data set consisted of immigrants and philopatric birds in a semi-isolated population in Sweden scored at 21 microsatellite loci. Fourteen cohorts were represented of which the four earliest were used to define a reference population. Female immigrants had lower assignment probability than males (i.e. were less likely to have been sampled in the reference population), and carried the majority of 'novel alleles' (i.e. alleles observed in the population for the first time). The difference in number of novel alleles between sexes was caused by a strong over-representation of females among the few individuals that carried several novel alleles, and there was a tendency for a corresponding female bias among individuals with low assignment probabilities. Immigrant males had similar or lower reproductive success than females. These results lead us to conclude that important interregional gene flow in great reed warblers depends on relatively few dispersing females, and that the novel allele technique may be a useful complement to the assignment technique when evaluating dispersal patterns from temporally structured data

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

Between-year variation of MHC allele frequencies in great reed warblers: selection or drift?

The major histocompatibility complex (MHC) genes are extremely polymorphic and this variation is assumed to be maintained by balancing selection. Cyclic interactions between pathogens and their hosts could generate such selection, and specific MHC alleles or heterozygosity at certain MHC loci have been shown to confer resistance against particular pathogens. Here we compare the temporal variation in allele frequencies of 23 MHC class I alleles with that of 23 neutral microsatellite markers in adult great reed warblers (a passerine bird) in nine successive cohorts. Overall, the MHC alleles showed a significantly higher variation in allele frequencies between cohorts than the microsatellite alleles, using a multi-variate genetic analysis (AMOVA). The frequency of two specific MHC alleles, A3e (P = 0.046) and B4b (P = 0.0018), varied more between cohorts than expected from random, whereas none of the microsatellite alleles showed fluctuations exceeding the expectation from stochastic variation. These results imply that the variation in MHC allele frequencies between cohorts is not a result of demographic events, but rather an effect of selection favouring different MHC alleles in different years

Partial albinism in a semi-isolated population of great reed warblers

Albinism in birds is thought to result from the expression of recessive alleles that disrupt melanin pigmentation at feather development. We have studied great reed warblers Acrocephalus arundinaceus in a recently founded and increasing population in Sweden during 15 years for the presence of birds with albinistic feathers. The study population was founded in 1978 and the few cases of albinism was exclusively recorded during the first five pears of our study (1985-1989). This fits to the expected pattern if albinism is governed by recessive alleles; we have previously demonstrated that the population suffered from inbreeding during the first years of our study. The albinistic birds experienced a similar lifetime reproductive success as normally coloured birds

Long-term maternal effect on offspring immune response in song sparrows Melospiza melodia

Knowledge of the causes of variation in host immunity to parasitic infection and the time-scales over which variation persists, is integral to predicting the evolutionary and epidemiological consequences of host–parasite interactions. It is clear that offspring immunity can be influenced by parental immune experience, for example, reflecting transfer of antibodies from mothers to young offspring. However, it is less clear whether such parental effects persist or have functional consequences over longer time-scales, linking a parent's previous immune experience to future immune responsiveness in fully grown offspring. We used free-living song sparrows (Melospiza melodia) to quantify long-term effects of parental immune experience on offspring immune response. We experimentally vaccinated parents with a novel antigen and tested whether parental vaccination influenced the humoral antibody response mounted by fully grown offspring hatched the following year. Parental vaccination did not influence offspring baseline antibody titres. However, offspring of vaccinated mothers mounted substantially stronger antibody responses than offspring of unvaccinated mothers. Antibody responses did not differ between offspring of vaccinated and unvaccinated fathers. These data demonstrate substantial long-term effects of maternal immune experience on the humoral immune response of fully grown offspring in free-living birds

Lemming–Food Plant Interactions, Density Effects, and Cyclic Dynamics on the Siberian Tundra

Theory predicts that trophic interactions can produce cyclic dynamics of microtine rodents, but that in addition, social interactions are necessary to create the cyclicity. We tested the induced defence hypothesis as a component driving multi-annual cycles by analyzing data on Siberian lemming (Lemmus sibiricus Kerr) populations and the levels of trypsin inhibitors (TI) and free proteins (SPP) in their food plants. We collected data at 12 sites along the Palaearctic tundra coast in 1994 and correlated these measures with lemming density and cycle phase. A negative correlation between lemming density and TI/SPP in Carex was found at the 12 sites. Cycle phase tended to correlate significantly with TI/SPP, and plants being grazed by lemmings respond with some delay. Mass of pancreas, the organ producing proteolytic enzymes, co-varied negatively with TI/SPP. These findings are in accord with predictions from the induced defence hypothesis. In contrast, reproductive effort did not conform to the predictions but co-varied strongly with density. Data are in accord with theory predicting that, in addition to trophic interactions, some self-regulation is necessary to create cyclic dynamics. Our data show that the plant defence hypothesis alone is not a sufficient explanation for the cyclicity. Several factors interact, including predation, food quality (including induced defence chemicals), and intra-specific competition.En théorie, les interactions trophiques peuvent produire une dynamique cyclique de rongeurs microtinés, mais en plus de cela, il faut des interactions sociales pour donner lieu à la cyclicité. Nous avons mis à l’épreuve l’hypothèse induite de défense en tant que composante intégrale des cycles multi-annuels en analysant les données relatives aux populations de lemming brun (Lemmus sibiricus Kerr) et les taux d’inhibiteurs de la trypsine (TI) et de protéines libres (SPP) se trouvant dans leur nourriture végétale. Nous avons recueilli des données à 12 emplacements le long de la côte paléarctique des toundras en 1994 et avons corrélé ces mesures à la densité des lemmings ainsi qu’à la phase du cycle. Une corrélation négative entre la densité des lemmings et TI/SPP dans Carex a été trouvée à 12 emplacements. La phase du cycle avait tendance à donner une corrélation significative avec TI/SPP, et les végétaux faisant l’objet d’un pâturage par les lemmings réagissaient avec un certain retard. Les masses de pancréas, soit l’organe producteur des enzymes protéolytiques, co-variait négativement avec TI/SPP. Ces constatations cadrent avec les prédictions émanant de l’hypothèse induite de défense. Par contre, les efforts de reproduction ne se conformaient pas aux prédictions mais co-variaient fortement en fonction de la densité. Les données sont en accord avec la théorie prédisant qu’en plus des interactions trophiques, il faut une certaine autorégulation pour créer une dynamique cyclique. Nos données montrent qu’à elle seule, l’hypothèse de défense des végétaux ne constitue pas une explication suffisante à la cyclicité. Plusieurs facteurs entrent en interaction, dont la prédation, la qualité de la nourriture (ce qui comprend les produits chimiques induits de la défense) et la compétition intraspécifique

Pollution related effects on immune function and stress in a free-living population of pied flycatcher Ficedula hypoleuca

We investigated whether exposure to heavy metal pollution affected the immune function of individuals in a free living population of a small insectivorous passerine bird, the pied flycatcher Ficedula hypoleuca. We measured humoral immune responses in two study areas: a polluted area in the vicinity of a copper smelter and a control area far from the smelter. Plasma corticosterone level and blood heterophil/lymphocyte ratio (H/L) were used as more general physiological measures of stress. The immune response of F hypoleuca was not suppressed by pollution stress. In contrast, we found that F hypoleuca males showed stronger Immoral immune responses to a novel antigen (tetanus toxoid) in the polluted environment than in the unpolluted one. After the immunization of males, numbers of lymphocytes rose significantly more in the polluted area, leading to a smaller H/L ratio than in males from the control area. Females showed no pollution related effects on their immune responses. Corticosterone levels of males and nestlings were not related to pollution levels. Nestlings showed somewhat higher H/L ratios and lower fledging success in the polluted area, both factors indicating increased stress levels in a polluted area. Our results suggest that Immoral immune response of male F hypoleuca may be enhanced under moderate levels of heavy metal Pollution. Enhanced immune function may, however, also be costly for birds and the higher humoral immune responses in polluted areas may thus have negative effects on the birds' breeding performance and survival