Estimating survival in continuous space from mark-dead-recovery data - Towards a continuous version of the multinomial dead recovery model.

Understanding spatially varying survival is crucial for understanding the ecology and evolution of migratory animals, which may ultimately help to conserve such species. We develop an approach to estimate an annual survival probability function varying continuously in geographic space, if the recovery probability is constant over space. This estimate is based on a density function over continuous geographic space and the discrete age at death obtained from dead recovery data. From the same density function, we obtain an estimate for animal distribution in space corrected for survival, i.e., migratory connectivity. This is possible, when migratory connectivity can be separated from recovery probability. In this article, we present the method how spatially and continuously varying survival and the migratory connectivity corrected for survival can be obtained, if a constant recovery probability can be assumed reasonably. The model is a stepping stone in developing a model allowing for disentangling spatially heterogeneous survival and migratory connectivity corrected for survival from a spatially heterogeneous recovery probability. We implement the method using kernel density estimates in the R-package CONSURE. Any other density estimation technique can be used as an alternative. In a simulation study, the estimators are unbiased but show edge effects in survival and migratory connectivity. Applying the method to a real-world data set of European robins Erithacus rubecula results in biologically reasonable continuous heat-maps for survival and migratory connectivity.

Estimation in the multinomial reencounter model - Where do migrating animals go and how do they survive in their destination area?

Spatial variation in survival has individual fitness consequences and influences population dynamics. Which space animals use during the annual cycle determines how they are affected by this spatial variability. Therefore, knowing spatial patterns of survival and space use is crucial to understand demography of migrating animals. Extracting information on survival and space use from observation data, in particular dead recovery data, requires explicitly identifying the observation process. We build a fully stochastic model for animals marked in populations of origin, which were found dead in spatially discrete destination areas. The model acts on the population level and includes parameters for use of space, survival and recovery probability. It is based on the division coefficient and the multinomial reencounter model. We use a likelihood-based approach, derive Restricted Maximum Likelihood-like estimates for all parameters and prove their existence and uniqueness. In a simulation study we demonstrate the performance of the model by using Bayesian estimators derived by the Markov chain Monte Carlo method. We obtain unbiased estimates for survival and recovery probability if the sample size is large enough. Moreover, we apply the model to real-world data of European robins Erithacus rubecula ringed at a stopover site. We obtain annual survival estimates for different spatially discrete non-breeding areas. Additionally, we can reproduce already known patterns of use of space for this species.

Age-specific reproduction in female pied flycatchers: evidence for asynchronous aging.

Age-related variation in reproductive performance is central for the understanding of population dynamics and evolutionary processes. Our understanding of age trajectories in vital rates has long been limited by the lack of distinction between patterns occurring within- and among-individuals, and by the lack of comparative studies of age trajectories among traits. Thus, it is poorly understood how sets of demographic traits change within individuals according to their age. Based on 40 years of monitoring, we investigated age-related variation in five reproductive traits in female pied flycatchers (Ficedula hypoleuca) including laying date, clutch size, brood size, nest success (probability that a nest produces at least one chick) and egg success of successful nests (proportion of eggs resulting in a chick). We disentangled within- from among-individual processes and assessed the relative contribution of within-individual age-specific changes and selective appearance and disappearance. Finally, we compared the aging pattern among these five reproductive traits. We found strong evidence for age-specific performance including both early-life improvement and late-life decline in all reproductive traits but the egg success. Furthermore, the aging patterns varied substantially among reproductive traits both for the age of peak performance and for the rates of early-life improvement and late-life decline. The results show that age trajectories observed at the population level (cross-sectional analysis) may substantially differ from those occurring at the individual level and illustrate the complexity of variation in aging patterns across traits.

Integrating stable isotopes, parasite, and ring-reencounter data to quantify migratory connectivity-A case study with Barn Swallows breeding in Switzerland, Germany, Sweden, and Finland.

Ecosystems around the world are connected by seasonal migration. The migrant animals themselves are influenced by migratory connectivity through effects on the individual and the population level. Measuring migratory connectivity is notoriously difficult due to the simple requirement of data conveying information about the nonbreeding distribution of many individuals from several breeding populations. Explicit integration of data derived from different methods increases the precision and the reliability of parameter estimates. We combine ring-reencounter, stable isotope, and blood parasite data of Barn Swallows Hirundo rustica in a single integrated model to estimate migratory connectivity for three large scale breeding populations across a latitudinal gradient from Central Europe to Scandinavia. To this end, we integrated a non-Markovian multistate mark-recovery model for the ring-reencounter data with normal and binomial mixture models for the stable isotope and parasite data. The integration of different data sources within a mark-recapture modeling framework enables the most precise quantification of migratory connectivity on the given broad spatial scale. The results show that northern-breeding populations and Southern Africa as well as southern-breeding populations and Western-Central Africa are more strongly connected through Barn Swallow migration than central European breeding populations with any of the African wintering areas. The nonbreeding distribution of Barn Swallows from central European breeding populations seems to be a mixture of those populations breeding further north and south, indicating a migratory divide.

Disruptive selection without genome-wide evolution across a migratory divide.

Transcontinental migration is a fascinating example of how animals can respond to climatic oscillation. Yet, quantitative data on fitness components are scarce, and the resulting population genetic consequences are poorly understood. Migratory divides, hybrid zones with a transition in migratory behaviour, provide a natural setting to investigate the micro-evolutionary dynamics induced by migration under sympatric conditions. Here, we studied the effects of migratory programme on survival, trait evolution and genome-wide patterns of population differentiation in a migratory divide of European barn swallows. We sampled a total of 824 individuals from both allopatric European populations wintering in central and southern Africa, respectively, along with two mixed populations from within the migratory divide. While most morphological characters varied by latitude consistent with Bergmann's rule, wing length co-varied with distance to wintering grounds. Survival data collected during a 5-year period provided strong evidence that this covariance is repeatedly generated by disruptive selection against intermediate phenotypes. Yet, selection-induced divergence did not translate into genome-wide genetic differentiation as assessed by microsatellites, mtDNA and >20 000 genome-wide SNP markers; nor did we find evidence of local genomic selection between migratory types. Among breeding populations, a single outlier locus mapped to the BUB1 gene with a role in mitotic and meiotic organization. Overall, this study provides evidence for an adaptive response to variation in migration behaviour continuously eroded by gene flow under current conditions of nonassortative mating. It supports the theoretical prediction that population differentiation is difficult to achieve under conditions of gene flow despite measurable disruptive selection.

West Nile virus monitoring of migratory and resident birds in Germany.

West Nile virus (WNV) is a mosquito-borne flavivirus naturally circulating in wild bird populations. The virus is also capable to infect a broad range of vertebrate species. Humans and equines are highly susceptible and can develop mild flu-like illnesses as well as severe encephalitis leading to fatalities. Most recently, WNV was found to circulate in countries close to Germany, such as France, Czech Republic, Italy, Austria, and Hungary. Given this epidemiological situation its spread to Germany cannot be ruled out. As no data on the WNV situation were available for Germany for the most recent past, we have conducted a serological survey to reveal WNV antibodies in wild birds. More than 2700 blood samples from migratory and resident birds representing 72 species that were collected during 2005-2009 were tested using an immunofluorescence assay and partly by micro-virus neutralization test. By immunofluorescence assay WNV-reactive antibodies could be demonstrated in 11 wild bird species. Similarly, WNV-neutralizing antibodies were revealed in migratory birds belonging to 10 species, but not in resident birds. According to the absence of WNV-reactive antibodies in resident birds and the absence of WNV-specific RNA in all investigated bird samples, there is currently no evidence for a WNV circulation in Germany.