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Genetics of Migration Max Planck Institute of Evolutionary Biology, Plön, Germany April 4th – 7th 2017 Tuesday 4th April 13.00 Arrival and registration 13.50 Welcome and overview of symposium Miriam Liedvogel, MPI Evolutionary Biology 14.00 15.00 KEYNOTE: Nancy Chen, University of California, Davis Deconstructing isolation-­‐by-­‐distance: the genomic consequences of limited dispersal 15.30 WORKSHOP: Quantitative Genetics Nancy Chen, University of California, Davis 18.00 Dinner and Discussion (buffet at MPI) Organisers Miriam Liedvogel (MPI Evolutionary Biology) Kira Delmore (MPI Evolutionary Biology) Christopher Jones (Rothamsted Research, UK) Refreshment break ____________________________________________________________________ Wednesday 5th April Session I: Insects as model to study migration/dispersal 09.00 KEYNOTE: Christine Merlin, Texas A&M University The molecular basis of monarch butterfly seasonal migration: a timing perspective. 09.45 10.05 10.25 Karl Wotton, University of Exeter, UK The environmental and genetic basis of migration in hoverflies Christopher Jones, Rothamsted Research, UK Insights into the flight potential of migratory moth pests from behavioural assays and transcriptomics Joan Ferrer Obiol, University of Barcelona, Spain Genome-­‐wide ddRAD sequence data based phylogeny of shearwaters, a group of pelagic seabirds exhibiting contrasting intra and interspecific migratory strategies 1 10.45 Refreshment break Session II: Genomics approaches to understanding architecture of migratory traits 11.15 KEYNOTE: Guojie Zhang, University of Copenhagen Bird genome 10K project, tracing the micro & macro evolutionary history of birds 12.00 12.20 12.40 Max Lundberg, Lund University, Sweden Genomic divergence across a migratory divide Kira Delmore, MPI Evolutionary Biology Demographic history and genetic basis of migration in the European blackcap Benjamin Van Doren, University of Oxford Integrated Zugunruhe programmes from migrants to residents: an annual cycle approach in Stonechats 13.00 Lunch (cafeteria) 14.00 WORKSHOP: CRISPR in Insects Christine Merlin, Texas A&M University 16.00 Refreshment break 16.30 FLASH TALKS 17.00 POSTER SESSION 18.00 Dinner and Discussion ____________________________________________________________________ Thursday 6th April Session III: Adaptation and Speciation 09.00 KEYNOTE: Claudia Bank, Gulbenkian Institute, Portugal Genetic incompatibilities in the presence of gene flow 09.45 Reto Burri, Friedrich Schiller University of Jena Linked selection in heterogeneous recombination landscapes: Prospects for the inference of adaptive evolution Amy Goldberg, Stanford University A mechanistic model for sex-­‐specific admixture Harald Ringbauer, Institute of Science and Technology Austria Inferring recent demography via isolation by distance of long shared genome blocks 10.05 10.25 2 10.45 Refreshment break 11.15 KEYNOTE: Jochen Wolf, Ludwig Maximilian University of Munich Speciation Genomics in Natural Populations 12.00 POSTER SESSION 13.00 Lunch (cafeteria) 14.00 WORKSHOP: Speciation Genomics Jochen Wolf, Ludwig Maximilian University of Munich 18.00 Dinner and Discussion ____________________________________________________________________ Friday 7th April Session IV: Expression profiling: Transcriptomics and Epigenetics 09.00 KEYNOTE: Melinda Baerwald, University of California, Davis Linking genes and the environment: differential DNA methylation in regulatory regions is associated with rainbow trout migratory-­‐related divergence 09.45 Clare Brown, Louisiana State University Comparative transcriptomics of avian migration: gene expression in the brains and livers of tree swallows (Tachycineta bicolor) staging for migration 10.05 10.25 Krista Nichols, NOAA Fisheries, Northwest Fisheries Science Center, Seattle Ontogenetic differences in gene expression in juvenile offspring of migratory and resident Oncorhynchus mykiss prior to the decision to migrate, mature, or remain resident Angelica Menchaca, University of Bristol, UK Genetic and morphological comparison of two populations of Tequila bats (Leptonycteris yerbabuenae) 10.45 Refreshment break 11.15 Roberto Frias Soler, Heidelberg University De novo annotation of the transcriptome and pre-­‐migratory gene expression changes of Northern wheatear, Oenanthe oenanthe (Aves) 11.35 11.55 Francisco Pulido, Complutense University of Madrid Measurement errors and genotype-­‐environment interactions – Why we need to consider them in molecular genetic studies of migration 12.15 Eirini Christodoulaki, University of Veterinary Medicine, Vienna Genomic signatures of migration and selection Closing remarks and end of symposium 3 Programme 4 Keynote Speaker Abstracts Nancy Chen University California, Davis http://nancy-­‐chen.weebly.com/ Deconstructing isolation-­‐by-­‐distance: the genomic consequences of limited dispersal Geographically limited dispersal can shape genetic population structure and result in a correlation between genetic and geographic distance, commonly called isolation-­‐by-­‐distance. Despite the prevalence of isolation-­‐by-­‐distance in nature, to date few studies have empirically demonstrated the processes by which it is generated within a pedigreed and genotyped population. Intensive, long-­‐
term demographic studies and exhaustive genomic surveys in the Florida Scrub-­‐Jay (Aphelocoma coerulescens) provide an excellent opportunity to investigate the influence of dispersal on genetic structure. Here, we use a panel of genome-­‐wide SNPs and extensive pedigree information to explore the role of limited dispersal in shaping patterns of isolation-­‐by-­‐distance in both sexes, and at an exceedingly fine spatial scale (within ~10 km). Isolation-­‐by-­‐distance patterns were stronger in male-­‐
male and male-­‐female comparisons than in female-­‐female comparisons, consistent with observed differences in dispersal propensity between the sexes. Using the pedigree, we demonstrated how various genealogical relationships contribute to fine-­‐scale isolation-­‐by-­‐distance. Simulations using field-­‐observed distributions of male and female natal dispersal distances showed good agreement with the distribution of geographic distances between breeding individuals of different pedigree relationship classes. Furthermore, we can accurately reconstruct observed isolation-­‐by-­‐distance patterns in autosomal and Z-­‐linked SNPs using coalescent simulations parameterized by the observed dispersal curve, population density, and immigration rate. Therefore, patterns of fine-­‐scale isolation-­‐by-­‐distance can be well understood as a result of limited dispersal over contemporary timescales in the Florida Scrub-­‐Jay. Christine Merlin Texas A&M University http://www.merlinlab.org/ The molecular basis of monarch butterfly seasonal migration: a timing perspective The eastern North American monarch butterfly (Danaus plexippus) has emerged as a powerful model system to study circadian clocks and their role in an unconventional output, the photoperiod-­‐
induced long-­‐distance migration. Circadian clocks are endogenous 24-­‐hour timekeepers that coordinate nearly all of the animal physiology and behavior to its environment to tune specific activities at the most advantageous time of the day. Monarchs use a circadian clock to navigate to their overwintering sites during their seasonal long-­‐distance migration. The clock time-­‐compensates for the movement of the sun across the sky over the course of the day and regulates the sun compass output in the brain. Circadian clocks may also be used to control the migratory switch and therefore regulate the genetic and epigenetic program controlling migratory physiology and behavior. I will discuss progress that our laboratory has made in developing reverse-­‐genetics approaches in the monarch butterfly to unlock its potential as a genetic model system to study animal clockwork mechanisms and the involvement of the circadian clock in seasonal migration 5 Guojie Zhang Section of Ecology and Evolution University of Copenhagen, Denmark China National Gene Bank BGI-­‐Shenzhen, China http://zhanggj.cngb.org/home/ Bird genome 10K project, tracing the micro&macro evolutionary history of birds Characterization of genomic biodiversity through comprehensive species sampling has the potential to change our understanding of evolution. To study evolution across a major vertebrate class, dissect the genomics of complex traits, and resolve a centuries-­‐old debate on the avian species tree, we formed a consortium focused on sequencing and analyses 48 bird genomes covered all 30 neognath orders, representing a wide range of avian evolutionary diversity. The phylogenomic analyses with full genome data produced a highly supported avian order phylogeny that resolves many debates on the timing and topology of their radiation. The new avian phylogenic tree has changed our view of the evolutionary process of many avian life-­‐history traits. Whole genome comparison for all bird species with other vertebrate species revealed several distinct macroevolution patterns of avian genome. The small genome size of bird was a consequence of massive loss of repeat elements and thousands of functional genes in bird ancestral stage. Different with other animal lineages that develop evolutionary innovation by obtaining new genetic materials, the bird genomes have extremely low new gene born rate. Instead, the bird genomes obtain millions of elements that specifically only conserved in bird lineages. These avian specific highly conserved elements are dominated by non-­‐coding regions and have significantly high level of enhancer potential. Through functional experiments, we demonstrate regulatory roles of ASHCEs in the punctuated evolution of dinosaur-­‐avian transition, and further highlight the importance of regulatory rewiring during macroevolutionary changes. In June 2015, we expanded the effort and have formally initiated Bird 10k genome project, which aims to sequence all the extant bird species in the world and reconstruct the tree of life for birds using the whole genome data. The B10K project will allow us to complete a genomic tree of life of the entire extant avian class, decode links between genetic variation and phenotype differences, uncover relationships of genetic evolutionary, biogeographical, and biodiversity patterns across a wide-­‐range of species, evaluate the impact of various ecological factors and humans on species evolution, and unveil the demographic history of an entire class of organisms Claudia Bank Gulbenkian Institute in Portugal https://evoldynamics.org/ Genetic incompatibilities in the presence of gene flow In this talk I will give an overview on recent work in the growing field of ‘speciation genomics’ unravelling the genetic underpinnings of adaptation and speciation from a micro-­‐evolutionary perspective. I will then present own work on a migratory divide in the barn swallow where evidence of divergent selection during migration and potentially also overwintering did not translate into genetic differentiation. Finally, I will discuss major challenges in interpreting patterns of genomic differentiation and their relationship to reproductive isolation using the Eurasian crow system as an example. 6 Jochen Wolf Ludwig Maximilian University of Munich http://www.ieg.uu.se/evolutionary-­‐biology/wolf/ Speciation Genomics in Natural Populations The view of species as entities amenable to evolutionary change elaborated by Charles Darwin laid the conceptual foundation for our current understanding of how biodiversity can be generated. Initially marred by a rudimental understanding of hereditary principles, evolutionists gained appreciation of the mechanistic underpinnings of adaptation and speciation following the merger of Mendelian genetic principles with Darwinian evolution. Spurred by a recent revolution in nano-­‐
sequencing technology speciation genetic research has become increasingly open to genetic non-­‐
model organisms. Genome-­‐wide processes can now be investigated at unprecedented resolution in essentially any eco-­‐evolutionary model species of interest. This development has expanded speciation research beyond the traditional boundaries and unveils the genetic basis of speciation from manifold perspectives and at various stages of the splitting process. Melinda Baerwald University of California, Davis http://gvl.ucdavis.edu/people/melinda-­‐baerwald/ Migration is essential for the reproduction and survival of many animals, yet little is understood about its underlying molecular mechanisms. We used the salmonid Oncorhynchus mykiss to gain mechanistic insight into smoltification, which is a morphological, physiological, and behavioral transition undertaken by some juveniles that culminates in a seaward migration. Given that gene x environment interactions and phenotypic plasticity are likely critical components to migratory divergence, epigenetic modifications (e.g., DNA methylation) may regulate gene expression and partially underlie migratory phenotypic diversity. To explore this, we quantitatively measured genome-­‐scale DNA methylation using reduced representation bisulfite sequencing of F2 siblings produced from a cross between steelhead (migratory) and rainbow trout (non-­‐migratory) lines. We identified 57 differentially methylated regions (DMRs) between steelhead and rainbow trout juveniles. DMRs were of high magnitude, ranging from 20-­‐62% differential methylation between life history types, and over half of the gene-­‐associated DMRs were in transcriptional regulatory regions. Many of the DMRs encode proteins with activity relevant to migratory-­‐related transitions (e.g. circadian rhythm pathway, nervous system development, protein kinase activity). This study indicates that differential DNA methylation at gene regulatory elements may be a critical molecular mechanism allowing interactions between an organism and its environment to balance phenotypic stability and plasticity, and ultimately modify migratory-­‐related phenotypes. 7 Speaker Abstracts Karl Wotton College of Life and Environmental Sciences, University of Exeter, Penryn Campus, Cornwall, UK [email protected] The environmental and genetic basis of migration in hoverflies My talk will focus on the migration of hoverflies, an economically important group of insects that provide dual ecosystem services of pollination and biological control of crop pests. Understanding the movement of these insects offers major opportunities to improve agricultural practices and to deepen our understanding of the environmental induction and genetic control of migration. I will discuss progress towards developing the hoverfly as a powerful new model for migration; one that will provide mechanistic insights into the function of migration-­‐associated genes. Christopher Jones Rothamsted Research, Harpenden, Hertfordshire, UK, [email protected] Insights into the flight potential of migratory moth pests from behavioural assays and transcriptomics Migration is a fundamental life-­‐history strategy for billions of insects including many of our most damaging pests. We still, however, know relatively little about the genes and biochemical pathways that drive this phenomenon in economically important insects. One obstacle to this is the challenge of quantifying migratory behaviour in the laboratory. Tethered flight assays allow flight performance to be measured in a controlled environment as a proxy for migration. We have recently begun to explore the applicability of these assays to quantify the variation in migratory flight in two major and global insect moth pests; the cotton bollworm (Helicoverpa armigera) and the fall armyworm (Spodoptera frugiperda). By flying adult moths from different populations and experimental treatments we show common patterns in flight performance across species with, for example, a strong positive correlation between flight duration and speed, indicative of persistent, straightened-­‐
out movement in those insects which fly furthest. We then use differences in flight performance between individuals to assign distinct phenotypes that can be used for downstream genetic analyses. One example of this is a transcriptomic study in H. armigera that reveals a suite of differentially expressed genes related to physiological adaptations required for long-­‐distance flight including the mobilisation of lipids as flight fuel, flight muscle structure and the regulation of hormones that influence migratory physiology. Extensions of this work to both epigenetic control and the relationship between migratory flight and disease will also be discussed. Joan Ferrer Obiol University of Barcelona, Spain [email protected] Genome-­‐wide ddRAD sequence data based phylogeny of shearwaters, a group of pelagic seabirds exhibiting contrasting intra and interspecific migratory strategies Bird species exhibit strikingly different scales and patterns of migration. A significant proportion of the phenotypic variance in migratory traits is genetic, but the genes involved in shaping these phenotypes are still largely unknown. Shearwater species exhibit contrasting intra and interspecific migratory strategies and thus, they provide an excellent model for investigating the genetic basis and evolution of different aspects of migratory behaviour such as migratory orientation, phenology or 8 duration. For that, it is instrumental to have well-­‐resolved phylogenetic relationships within the group. Nonetheless, understanding their systematics has been challenging. Previous phylogenetic studies using fine-­‐scale morphological details, osteological characters and single mitochondrial gene sequences have provided poor resolution possibly due to their complex and overlapping morphological variation, incomplete lineage sorting and speciation with gene flow. To clarify these relationships, we are generating double-­‐digest restriction site-­‐associated DNA sequencing data (ddRADseq) for every species and subspecies of shearwater with the aim of building a robust phylogeny of the group and to test for gene flow during the diversification of the genus. In a first step, we obtained an average of 38,837 ddRAD-­‐tag loci per sample. The number of homologous loci shared among samples was negatively correlated with genetic distance, probably due to the accumulation of restriction site polymorphisms. Nonetheless, for all samples combined, we obtained 2,024 homologous loci despite ~14 my divergence among the three shearwater genera (Ardenna, Calonectris and Puffinus). Phylogenetic analyses of concatenated data and species tree methods generated broadly consistent results. Species tree methods allowed to visualise uncertainty in the majority-­‐rule topology and introgression tests, specifically designed to detect ancient gene flow (e.g. ABBA/BABA tests), allowed to test the role of gene flow during speciation in these pelagic seabirds. The clear resolution of this study will establish a framework for investigating the genetic basis of migratory traits in shearwaters. Max Lundberg Department of Biology, Biologihuset, Lund University, Sweden [email protected] Genomic divergence across a migratory divide Migration requires adaptations in behavioural, morphological and physiological traits. These traits have been shown to have a strong genetic basis, but very little is known about the underlying genes. Here we use whole-­‐genome resequencing and a customized snp array to detect genetic differences between two differentially migrating subspecies of the willow warbler Phylloscopus trochilus. We find that the genomes are extremely similar, with differences clustered in three divergent chromosome regions that potentially represent inversion polymorphisms. One of the regions does not contain alleles that are strictly associated with each of the subspecies, but instead appears to be associated with adaptation to high altitude and latitude. The other two chromosome regions contain alleles specific to either subspecies and may contain differences related to the different migratory phenotypes of the subspecies. Kira Delmore Max Planck Institute for Evolutionary Biology, Plön, Germany [email protected] Demographic history and genetic basis of migration in the European blackcap The amazing accuracy of migratory orientation across the animal kingdom is facilitated by the use of magnetic and celestial compass systems that provide individuals with both directional and positional information. Quantitative genetics analysis of several animal systems suggests that migratory orientation has a strong genetic component. Nevertheless, the exact identity of genes controlling orientation remains largely unknown, making it difficult to obtain an accurate understanding of this fascinating behavior on the molecular level. The European blackcap is a songbird that breeds in Central Europe. This species exhibits substantial variation in migratory orientation, with populations migrating northwest, southwest and southeast. We have capitalized on this variation to narrow in on the genes that regulate migratory orientation. Specifically, we have employed state-­‐of-­‐the-­‐art next 9 generation sequencing techniques to assemble a draft reference genome for this species, examine their demographic history and identify genomic differences between populations with different migratory orientations. We find that the ancestral state within this group is resident, evidence that these resident populations were not affected by the ice sheets and identify small regions of the genome that likely code for differences in migration. We discuss within these regions in terms of their function and ontology. Our results relate not only to which genes control migratory orientation but also when variation for this trait is encoded and how it translates into differences in behaviour, with important implications for understanding the ecology and evolution of seasonal migration. Benjamin Van Doren Edward Grey Institute of Field Ornithology, Department of Zoology, University of Oxford, Oxford, UK [email protected] Integrated Zugunruhe programmes from migrants to residents: an annual cycle approach in Stonechats Studies of captive birds have revealed an inherited programme for Zugunruhe— “migratory restlessness” — and key features of migratory timing and navigation. Zugunruhe is part of a more comprehensive annual-­‐cycle programme, which coordinates fluctuations in many aspects of physiology, for example in the immune and metabolic systems. In a suite of studies, we have compared annual cycles of populations and hybrids of stonechats (genus Saxicola) with diverse migratory phenotypes. The overarching picture is that annual-­‐cycle organisation is generally conserved in stonechats, but shows endogenous, population-­‐specific modifications. In the most migratory population, behaviour and physiology showed the greatest intra-­‐annual fluctuations, for example of immune regulation, and at least for Zugunruhe, the greatest inter-­‐annual stability (for example, despite increasing age). Hybrids generally showed intermediate patterns, but in some traits resembled one of the parental groups. These findings generally confirm ideas that evolution of a “migratory syndrome” has involved an integrated, gradual change in multiple traits including Zugunruhe. Hence, rewarding strategies for identifying the genetic basis of migration should look at multiple integrated genomic changes and at modified seasonal dynamics of gene expression and epigenetic regulation in corresponding traits. As a first step towards investigating the genetic basis of migratory phenotypes among stonechats, we have established patterns of genetic diversity and differentiation by whole-­‐genome pooled sequencing in stonechats from five populations. Reto Burri Friedrich Schiller University of Jena reto.burri@uni-­‐jena.de Linked selection in heterogeneous recombination landscapes: Prospects for the inference of adaptive evolution Genome-­‐scale data empower genome-­‐wide perspectives on adaptive evolution. However, the identification of genome regions underlying contemporary adaptation is complicated by diverse evolutionary processes that obscure and mimic the footprints of positive selection. Particularly the long-­‐term effects of linked selection in heterogeneous recombination landscapes are difficult to account for and yet often poorly acknowledged. I showcase how this process affects the distribution of genetic diversity along genomes. Based on recent research on the evolution of genome differentiation across the speciation continuum I then argue that the interpretation of differentiation landscapes depends on the time scales studied, and that a comparative population genomic approach can help overcome shortcomings of traditional genome scans for adaptive evolution in extant populations and species. 10 Amy Goldberg Stanford University, California, USA [email protected] A mechanistic model for sex-­‐specific admixture Animal populations often experience sex-­‐biased dispersal and admixture, owing to factors such as sex-­‐specific breeding pressures or inbreeding avoidance. These processes have been empirically studied through comparisons of genetic ancestry for the X chromosome, which is inherited sex-­‐
specifically, and that of the autosomes, which are not. Using a mechanistic admixture model, we derive recursive expressions for the time-­‐dependent behavior of X-­‐-­‐chromosomal ancestry for general migration and admixture histories. For a model with constant contributions over time, we provide analytical results for the mean X-­‐-­‐chromosomal ancestry component, comparing ancestry on the female and male X chromosomes in an admixed population to corresponding values on the autosomes. Using this framework, we consider the sex-­‐specific history of two major migrations in European prehistory: the spread of agriculture to Europe from Anatolia during the early Neolithic and the late Neolithic/Bronze Age migration from the Pontic-­‐Caspian steppe. Contrary to previous hypotheses suggested by the patrilocality of many agricultural populations, we find no evidence of sex-­‐biased admixture during the migration that spread farming across Europe. For later migrations from the Pontic steppe, however, we estimate a dramatic male bias, with ~5-­‐14 migrating males for every migrating female. Harald Ringbauer Institute of Science and Technology Austria, Klosterneuburg, Austria [email protected] Inferring recent demography via isolation by distance of long shared genome blocks Recently it has become feasible to detect long blocks of genome shared between pairs of samples. These blocks are direct traces of recent coalescence events and as such they contain ample signal for inferring recent demography. Here we examine sharing of such blocks in two-­‐dimensional populations with local migration. Intuitively, the chance of co-­‐inheriting long blocks decreases with growing pairwise distance. Using a general diffusion approximation, we derive formulas for this isolation by distance pattern of long IBD-­‐blocks and show that mean block sharing quickly approaches exponential decay for growing sample distance. An inference scheme based on our results can robustly estimate the dispersal rate and effective population density, which we demonstrate on simulated data. As an example, we apply it to Eastern European samples in the POPRES data set to estimate mean migration distances and the rate of recent population growth. Clare Brown Department of Biological Sciences and Museum of Natural Science, Louisiana State University, USA [email protected] Comparative transcriptomics of avian migration: gene expression in the brains and livers of tree swallows (Tachycineta bicolor) staging for migration Avian migration has been a subject of scientific inquiry for over two hundred years, and ornithologists have amassed a substantial body of knowledge on the ecology, physiology, geography, and evolution of the seasonal movements of birds. One rapidly advancing area is the use of high-­‐
throughput sequencing and bioinformatics to study migration and migratory species. These techniques can be applied to the question of which genes and genomic regions are involved in migration. We used a comparative transcriptomic approach to search for candidate genes for 11 migration in Tree Swallows (Tachycineta bicolor), a long distance migrant that breeds across North America and winters along the Gulf Coast, and in Florida, Central America, and the Caribbean. In late fall, enormous roosts of Tree Swallows form in agricultural fields in the southeastern United States. These roosts are believed to be migratory stopover sites. There is substantial variation in physical condition (e.g. furcular fat) of birds captured in these roosts, suggesting corresponding variation in migratory schedule. This variation in migratory timing enabled us to compare gene expression in brain and liver among thirty individual swallows captured at the same time, under the same environmental conditions, at a roost near Vacherie, Louisiana. The captured individuals fell into three furcular-­‐fat classes, suggesting that these birds can be grouped into three categories of readiness to migrate. We extracted RNA from the brains and livers of these birds, and sequenced their whole-­‐brain and liver transcriptomes with Illumina HiSeq. We developed a novel, reference-­‐
transcriptome free method to identify differentially expressed transcripts among the three fat classes. These differentially expressed transcripts can be considered as candidate genes for involvement in migratory behavior and physiology. Krista Nichols NOAA Fisheries, Northwest Fisheries Science Center, Seattle, Washington State, USA [email protected] Ontogenetic differences in gene expression in juvenile offspring of migratory and resident Oncorhynchus mykiss prior to the decision to migrate, mature, or remain resident Residency and anadromy represent the extremes of life history variation within salmonid fishes, but the genetic underpinnings of life history variation are still poorly understood, both with regard to the genes involved in differentiation and the timing of the onset of differentiation. We examined differential gene expression in offspring from migratory and resident O. mykiss, to better understand the genes and molecular processes potentially involved in the process of life history determination, and when in development the differentiation is initiated. RNA sequencing was conducted in O. mykiss from Sashin Creek and Sashin Lake, Alaska. Families were reared in a common environment and brains were sampled at 4-­‐6 month intervals from hatch until June of their second year when smoltification or precocious maturation occurs in this system. A reference transcriptome was created and differential expression analysis was performed including 6 time points. Differential gene expression between migratory and resident offspring was observed as early as hatch and throughout development, and was highly dependent on both time and sex. Enrichment of differentially expressed genes was identified for a number of biological pathways, including phototransduction pathways, which can regulate circadian rhythms, ultimately leading to the decision to migrate or remain resident. Angelica Menchaca School of Biological Sciences, University of Bristol, UK [email protected] Genetic and morphological comparison of two populations of Tequila bats (Leptonycteris yerbabuenae) The tequila bat (Leptonycteris yerbabuenae) is a long distance migratory species that pollinates plants of economic importance. The species subdivides in two populations where females travel in the spring from central Mexico to southern US covering a distance of over 1400km and give birth in early summer; a second population remains in the south and gives birth in late autumn. Mating is believed to occur in the hibernacula and it is possible that individuals from both populations share these roosts with the males. Analysis of genetic substructure show high levels of gene flow between 12 populations, however, colonies differ in their migratory behaviour. Here, we compare blood and brain transcriptomes of individuals at similar reproductive state to evaluate differences in gene expression and potentially identify genes and pathways in the migratory behaviour of the species. In addition, morphological differences between these two populations were evaluated with geometric morphometrics to assess variation in wing shape and compare flight efficiency. Roberto Frias Soler Heidelberg University, Germany [email protected]­‐heidelberg.de De novo annotation of the transcriptome and pre-­‐migratory gene expression changes of Northern wheatear, Oenanthe oenanthe (Aves) The flying abilities and especially long migrations are between the most outstanding features in birds. They involve enormous metabolic, physiologic, and morphological challenges, not all of them well known at genetic level. In this work we present the partial transcriptome of the small migrant passerine Northern wheatear (Oenanthe oenanthe), by using RNA-­‐Seq. The transcriptome was constructed de novo from six organs: Brain, Muscle, Intestine, Liver, Adipose tissue and Skin. All individuals were captive birds, and the samples were collected at three different points of time before the winter migration. There were identified 14724 genes (17539 isogroups and 21982 isotigs), 37 % of them with less than 50 % identity with any other gene from turkey and zebra finch genomes or the Swissprot and NCBI databases. Around 13 % of the isogroups have a putative splicing variant, most of them just a second alternative. The biological processes better represented are related with ATP production, lipid and glucose metabolism, and muscle structure. In each tissue in between 200 to 2000 genes were significantly regulated among three different sampling points. Francisco Pulido Department of Zoology and Physical Anthropology, Complutense University of Madrid [email protected] Measurement errors and genotype-­‐environment interactions – Why we need to consider them in molecular genetic studies of migration For conducting genetic studies of behaviours, it is essential to clearly define the behaviours under study and to measure them accurately and with precision. Low repeatabilities, due to measurement errors, will result in low heritabilities and low statistical power when trying to detect potential genetic effects on these behaviors. Migratory traits are often difficult to define, as they are often shared with non-­‐migratory individuals, and difficult to measure, as they are expressed by moving animals in the wild over an extended period of time. An additional problem in the genetic study of migratory traits is that they are highly flexible, both within and among individuals, and that their expression often depends on environmental conditions. To account for the condition-­‐dependent expression of many migratory traits, it is necessary that environmental conditions are standardized or randomized when migratory behaviour is measured and that potential genotype-­‐by-­‐environment effects are taken into account. The environmental threshold model of migration predicts that this problem will be particularly important in populations that are close to the migration threshold, i.e. in partial and short-­‐distance migrants, as we find, for instance, in Spanish blackcaps. A recent study on the association between candidate genes and migratory behavior yielded different results, depending on the level (e.g. within or among populations), the trait analyzed and the environmental conditions animals experienced. These results suggest that for studying the genetics of migration in these populations we need complex experimental designs to separate environmental and genetic effects. Generally, any molecular genetic study of migratory behaviour should clearly define and 13 describe the behavioral components analyzed, calculate and optimize measurement repeatability and measure and control for the effects of environmental conditions on its expression. Eirini Christodoulaki Institute of Population Genetics, University of Veterinary Medicine, Vienna, Austria [email protected] Genomic signatures of migration and selection Polymorphic populations that evolved for many generations in an environment under specific conditions have most likely adapted to these environmental conditions by changing allele frequencies at loci contributing to a higher fitness. Migrating these evolved genotypes into a population that evolved in different environmental conditions allows to test the behaviour of alleles beneficial in the previous environment. The fate of these alleles in the recipient population depends on the strength and direction of selection (s) that acts on them, the proportion of genotypes that migrate every generation (m) and the size of the population (N). Although there is well established theory, empirical studies that could be combined with theoretical expectations are rare. In this project we are interested in the fate of the beneficial alleles when they migrate into a non-­‐adapted population and also, the fate of deleterious alleles when they migrate into a well-­‐adapted population. We use populations of D. simulans that evolved in different environments and we try to unravel the genomic signatures of migration between differentially adapted populations. 14 Poster Abstracts Elinor Jax Max Planck Institute for Ornithology [email protected] Genetic diversity and evolution of the toll-­‐like receptor signaling pathway in mallards (Anas platyrhynchos) Large population sizes and short generation times of pathogens are believed to create and retain high genetic variation of the immune system of host populations. Animals that migrate between different environments further encounter different pathogen communities during their lifecycle, and this might influence their immunogenetic diversity. Whilst allelic variation and its consequences have been intensively studied in the adaptive immune system, such as the major histocompatibility complex (MHC) in birds, the extent and effect of genetic variation in the innate immune system is not well understood. In this study we investigated the genetic variation in a group of innate immune genes in four populations of wild mallards (Anas platyrhynchos), and evaluated and compared the evolutionary patterns of these genes. The mallard is a great model system for eco-­‐immunology studies. It successfully inhabits all continents (except Antarctica) and therefore allows comparisons of immunocompetence in populations inhabiting a wide range of environments, and differing in important life history traits such as migration strategies. We used hybrid capture and next generation sequencing to genotype 142 immune genes from the toll-­‐like receptor signalling pathway. We specifically assessed whether innate immune genes are spatially structured, to evaluate whether the innate immune system is adapted to local pathogen communities. We further studied selection patterns in the genes across the pathway to improve our knowledge of the evolution of the innate immune system in birds. We discuss our results in the context of disease ecology. Teja Curk NIOO-­‐KNAW Netherlands Institute of Ecology [email protected] Revealing migratory pathways and changes in migration behaviour of Eurasian blackcaps using multi-­‐state models Bird species are challenged to adapt to rapidly changing environmental conditions by adjusting their migratory strategies. Several studies in the past decades have shown that Eurasian blackcaps are capable of adjusting their migratory habits in just a few generations. It is generally believed that since 1960 an increasing part of the Central European blackcap population is wintering in the British Isles rather than in the western Mediterranean. However, direct evidence for this phenomenon is still lacking, since the idea for this shift in migratory routes is entirely based on orientation experiments with captive blackcaps coupled to observations of an increased wintering population of blackcaps in the British Isles. Furthermore, changes in migration routes might have evolved also in other populations in Europe that we have no current knowledge about. In this study we use all available ring-­‐reencounter data of blackcaps to investigate their migratory pathways and potential changes that occurred through time. A migratory connectivity model is used to account for spatial variation in re-­‐encounter probability which represents a major bias in ring-­‐reencounter data. The model results showed that populations east from the migratory divide, well visible in Austria and Czech Republic, in autumn mainly migrate SE whereas populations west from the divide mainly migrate SW. Migrants from continental Europe and the breeding population from the British Isles do not in large proportions winter in the British Isles. The enormous number of British wintering blackcaps can rather be explained by the overall increase in the blackcap population in Europe. Since 15 1980, residency seems to have increased and migration decreased over much of Europe. Reduced migration most likely evolved as an adaptation to climate change. Our study provides a step forward towards understanding migratory pathways of blackcaps. Hannah Justen, Timo Hasselmann University of Kiel, University of Lübeck, MPI Evolutionary Biology, Germany [email protected], [email protected]­‐luebeck.de Characterisation of Clock gene polymorphism across Saxicola populations and migratory phenotypes Timing is essential for the survival and successful reproduction of migratory birds. The Clock gene has been identified as candidate gene for circannual timing processes, it is extremely homologous except for one variable exonic region that shows poly-­‐Q lengths variation. Longer repeats have been associated with higher breeding latitude for various bird species. Some studies have also suggested a link of Clock gene variability with migratory timing. Here we characterise Clock gene variability across different stonechat populations at varying breeding latitude that also exhibit clear differences in migratory behaviour. The Saxicola complex allows us to examine genotype differences in closely related populations with different phenotypes, including one (resident) African population breeding at the equator experiencing the same 12:12 light/dark cycle throughout the year. Interestingly, the African stonechats showed little variability in clock genotype, while more variation was detected in the other populations, suggesting that the lack of light-­‐entrainment might have selected for an optimized Clock genotype. The most variable Clock genotype also including longest alleles was found in Siberian stonechats, supporting previous findings with longer repeats found at higher breeding latitudes. Evaluating Clock gene variability in the context of migratory phenotype shows a stronger signal in autumn compared to spring, with shorter alleles associated with earlier timing of autumn migration. Alexandre Lemopoulos University of Eastern Finland [email protected] Genetic components in the migration behaviour of brown trout Of all the migrating animals, salmonids fishes are among the most famous and economically important ones. As many salmonids, brown trout present two different type of life history-­‐thus two morphs-­‐ linked to migration. The first one is a migrating morph, where trout leave from rivers to join either the sea (anadromous sea trout) or large lakes (adfluvial trout). The second one is a resident morph where the fishes stay in flowing rivers for their entire life. The reasons that lead to these different life history traits are still uncertain and are thought to arise out of several origins. The heritability of the migration potential is well documented, indicating a genetic influence over this trait. Yet, no studies have been able to single out a clear explanation for these different life history strategies. In this study we focus on the implication of genetics in the migrations of brown trout. We used next-­‐generation sequencing of RAD tags and obtained 3’390 SNPS. We compared 4 Finnish watersheds with both migrant and resident individuals. For each population, 30 individuals were sampled and sequenced. By comparing migratory and resident trout populations, we were looking for genomic regions associated with the different migratory patterns and indicating differences in life history traits. So far we have identified 125 outlier SNP. Our aim is to annotate them and understand whether they could potentially be involved in the migration behaviour. A better understanding of the migrations processes could lead to major advances in the managing and the conservation of brown trout and its stocking all over the world. 16 Gillian Durieux, Juan Sebastian Lugo Ramos Max Planck Institute for Evolutionary Biology, Plön [email protected], [email protected] Dissecting the Neurogenomics of European blackcap (Sylvia atricapilla) migration The migratory behaviour of European blackcaps has received much attention in recent decades. Classic works from Peter Berthold, Andreas Helbig and others, have demonstrated that both migratory distance and orientation have heritable components. Nonetheless, the identity of the elements or mechanisms that are inherited remain elusive. Recent population genomics approaches allow us to de novo identify potential candidates that might play a role in migratory behaviour. But even small differences on the sequence level might result in big differences on the gene expression level, and different tuning of signaling cascades might shape variability in migratory phenotype. We are combining RNA-­‐seq analyses with ChIP-­‐seq analyses to obtain gene expression profiles and characterize epigenetic patterns in key brain areas thought to be involved in regulating migratory behaviour: the hippocampus, Cluster N and the Suprachiasmatic Nuclei of the brain. In addition to comparing populations with differing orientation strategies from either side of a migratory divide, we also make use of the fact that the migratory behavior is only expressed during the migratory season. Thus we contrast gene expression profiles of both populations (i) during migration, and (ii) contrast this pattern with expression patterns obtained during a control period (i.e. outside the migratory season). To characterize the birds’ activity profile in order to identify control behavior versus migratory restlessness behaviour we monitored their day and night activity, using infrared motion loggers and IR camera observation. With this approach we intend to have a clear image of the neuroanatomical and expression dynamics elements playing a role in migratory behaviour. Vaishnovi Sekar Lund University, Sweden & University of Konstanz, Germany [email protected] Genome-­‐wide patterns of immune related genes across differently migrating goose species The mallard (Anas platyrhynchos) has been established as the main host of the Avian Influenza Virus (AIV) and has the best understood immune system among its relatives: ducks, the geese and swans (family Anatidae). Anatidae inhabit aquatic habitats and are therefore exposed to a high prevalence of diseases because wetlands are ideal ecosystems in which viruses and bacteria can survive for a prolonged time. Yet it is known from field observations and experimental evidence that some groups within the Anatidae are more often or more heavily infected by some diseases, for example, AIV. While ducks are mostly carriers of such viruses but do not suffer disease, geese and swans are often reported to show high mortality in disease outbreaks of certain strains. True geese (Anserini) are closely related to the mallard (Anas platyrhynchos) but are more susceptible to AIV. However, the genetic basis of this is not known. It is also evident that pathogen fauna may differ with geography and thereby migratory distance and species range will have impact on the immunity of these birds. Geese are therefore an interesting group for discovering what differences in the genome might be associated with the differences in susceptibility to pathogens among the waterfowl. Recently, the genomes of all geese have been re-­‐sequenced and subsequently their phylogenetic relationships were resolved. Beyond this taxonomic assessment, further investigations on the specific genes involved in immunity have not been performed yet. In this study, we investigated the genetic diversity in immune genes responsible for responses to avian diseases across all goose species. We performed reference-­‐based annotation to predict immune genes in the goose genomes by comparative mapping to the mallard genome and transcriptome, as well as by gene prediction tools such as MAKER. With the sequence information across a wide range of immune genes, we analysed 17 the distribution of immunogenetic diversity and signatures of natural selection among all geese in many genes. Our comparisons focus on different life histories relating to migratory lifestyles of the goose species. Jesko Partecke Max Planck Institute for Ornithology [email protected] Quantifying reactions norms of songbird migration in Euroasian blackbirds We live in a golden era for studies of animal movements in the wild. Technological advances allow the precise lifetime tracking of individuals around the globe. We can now finally test environmental influences on behavioural traits of natural populations by experimentally manipulating these traits, selecting for specific traits in captivity and releasing these individuals back into the wild to test their reaction norms in relation to changing environments. We study Eurasian blackbirds because they are partially migratory in Germany, entirely migratory in Russia, and stationary year-­‐round in France. We first observe individual birds in the wild and determine their natural migration behaviour in highest resolution. Second, we breed individuals in our large (common garden) aviary facilities. We then release captive bred individuals back into each of the three parent environments. We will also selectively breed lines of migratory and non-­‐
migratory blackbirds to test whether they decide to migrate or not when released into the wild, based on their genetic background and the current environmental conditions. This study will determine how flexible the reaction norm of migration behaviour is. We concurrently study the genomic and physiological background of individuals differing in migration strategies to link genomic bases and energy metabolism to migration decisions. Peter Ranke Norwegian University for Science and Technology [email protected] Demographic consequences of translocation: an experiment in a house sparrow population In conservation biology translocation is a powerful tool in augmenting small-­‐ and threatened populations. We examine the sex-­‐specific differences in reproductive success and survival between translocated and resident individuals in a merged population of house sparrows Passer domesticus. From individual data we estimate the contribution from the translocated individuals to the growth of the total population. We found evidence for assortative distribution of parentages based on origin: there were fewer offspring between translocated males and resident females than expected, and hence fewer fledglings produced by mixed pairs. Interestingly, the reproductive success of translocated males depended on the throat badge size, and only translocated males with large badge size were as successful as resident males. However, offspring from pairs with mixed origin had higher recruitment probability, which suggest hybrid vigour, alleviating the negative effects of being translocated. The contribution from translocated and resident individuals to the stochastic component of the long-­‐run growth rate of the population was therefore similar; neither the mean individual contribution in fitness nor the demographic variance differed between the two groups. Thus, this experiment shows that translocated individuals may have a similar demographic influence on the growth of local populations as resident individuals. Still, the intermixing of individuals with different origin may be low, and fitness may differ according to origin in relation to individual differences in sexually selected traits. Additionally, in our experiment hybrid vigour seems to have relaxed the negative reproductive consequences of assortative distribution of parentages to some extent. 18 Jaspar van Heusden Universidad Complutense Madrid [email protected] Association between candidate gene loci and migratory behaviour in Iberian blackcaps (Sylvia atricapilla) Heritable components have been identified as major causes of differences in the expression of specific migratory behaviour, yet our knowledge of its molecular bases remains limited. A first study on blackcaps (Sylvia atricapilla) sampled over its entire range demonstrated an association both between among-­‐population and within-­‐population variation in migratory behaviour/nocturnal restlessness and mean allele length on the candidate gene locus ADCYAP1. Studies on other species could not fully replicate this result. We aimed at investigating the association between candidate loci and migratory behaviour in the blackcap on a smaller geographic scale to assess the generality of this association. We genotyped 564 birds from 21 Iberian populations using 13 microsatellite loci, 4 of which were candidate gene loci (ADCYAP1, CLOCK, NPAS2 and SERT). The results confirm previous findings that on average longer alleles at the ADCYAP1 locus are found in the more migratory populations. In addition to the results from the study in blackcaps at a continental scale, we found an association between allelic variation at the CLOCK gene with variation in migratory behaviour on the Iberian Peninsula. In a second study, we kept 104 birds, collected as fledglings, from 3 of the previously studied populations (a migratory, a partially migratory and a sedentary population) in captivity and tested whether migratory activity in captivity (i.e. migratory restlessness) reflects migration behaviour in the wild. We tested for correlations between migratory restlessness and the same set of genetic markers as before. In this sample, migratory restlessness was not correlated with any genetic marker. We further found that among population differences in migratory behaviour in Iberian blackcaps, as they were found in the wild, are not expressed under our laboratory conditions. This unexpected result could be due to environment-­‐dependent thresholds in birds from different populations. These result suggest that conclusions on the genetics of migration derived from studies in the laboratory may be misleading if genotype-­‐by-­‐environment interactions are not accounted for. Bernhard Voelkl University of Bern, Swutzerland [email protected] Are changes in migration habits in the northern bald ibis associated with changes in DNA methylation patterns? Due to changes in climate, resource and habitat availability migratory pattern change considerably and new migration patterns have been observed emerging and other formerly migrating populations ceased to migrate. The fast rate at which these changes manifest themselves in the populations suggests a rapid evolutionary change in key regulatory mechanisms. Recent findings suggest that changes in chromatin marks-­‐-­‐most prominently in DNA methylation patterns-­‐-­‐can be an efficient mechanism for incorporating information about environmental changes into the hereditary system. Given the frequent and rapid changes in migration habits we suggest that these changes could be governed by changes in chromatin marks. To investigate this we suggest an exploratory study in the course of a re-­‐introduction program of Northern bald ibis (Geronticus eremita) in central Europe (LIFE+ Northern Bald Ibis). For this re-­‐introduction scheme eggs from non-­‐migrating birds from zoo stocks are taken over by human foster parents for hand rearing. At an age of four months these birds will be guided by their foster parents over about 1000 km to their designated wintering area in Tuscany using ultralight airplanes. At their destination the birds are finally released and they will start returning to their breeding grounds north of the alps after reaching sexual maturity. We aim at comparing DNA samples from birds at different stages: From the hand raised birds before and after 19 their first migration, from siblings or half-­‐siblings of those birds which are not migrating, from the parent generation (non-­‐migrating) and from the next generation of migrating birds with migrating parents. Taken together this will allow us making both cross-­‐sectional and trans-­‐generational comparisons. Angela Parody-­‐Merino Massey University, New Zealand [email protected] Associating genetic variation with migration departure time variation in a long-­‐distance migratory bird Describing mechanistic links between genetic variation and behaviour level variation in free-­‐living animals presents a major scientific challenge. Migratory bar-­‐tailed godwits (Limosa lapponica) that breed in Alaska and over-­‐winter in Australasia (subspecies baueri) perform the longest recorded avian non-­‐stop migratory flight, directly from Alaska to New Zealand (N.Z.), c. 12,000 km away. Individuals migrate back to Alaska via a refuelling stop-­‐over in eastern Asia, leaving N.Z. across an approximately 5-­‐week time period from late February to early April. Individual godwits display remarkable year-­‐to-­‐year consistency in when they leave N.Z. The control of avian migration timing is believed to involve changes in daylength as perceived by the birds, but this insight does not explain the observed inter-­‐individual variation in different departure schedules. It is anticipated that genetic variation is causally and mechanistically associated with such phenotypic variation. To investigate this possibility, we are looking for associations between individual migration departure times and genotypic variation at loci that form elements of the endogenous circadian clock, and its outputs, and components of the Hypothalamic-­‐Pituitary-­‐Gonadal (HPG) axis. A reference bar-­‐tailed godwit genome has been assembled and annotated. SNPs considered to merit genotyping are being identified after re-­‐sequencing and comparison of 19 additional genomes from 9 extremely early-­‐
departing and 10 extremely late-­‐departing individuals. Once a manageable number of SNPs in plausible loci have been selected we will genotype these in the genomes of 165 individuals with well-­‐
established migration departure date phenotypes. It is hoped that the results of this study will provide insights into the mechanisms underlying vertebrate migration timing and vertebrate seasonality in general. Gregor Schalm University of Oldenburg gregor.schalm@uni-­‐oldenburg.de Dispersal duration may be associated with PolyQ-­‐length in damselfish The life history of most coral reef fish is characterised by a dispersal phase, during which larvae leave their natal reef after hatching for an often weeks-­‐long dispersal in the open ocean. Pelagic larvae duration is species depending and differs in damselfish (Pomacentridae) between 4 and 34 days post hatching; this duration influences the geographic distribution of coral reef fish. Only a single non-­‐
dispersing species is known, in which both parents defend the brood against high predation pressure. To understand the underpinning genetic basis of dispersal duration, we sequenced and compared a part of the gene clock (the PolyQ-­‐allele) with strong length polymorphisms in 11 different species of the pomacentrid family. Interestingly, the preliminary data show a correlation between the pelagic larval duration, and the number of glutamine in the studied clock sequence for fish caught at the Great Barrier Reef, Australia. It is a first hint, for PolyQ being an important factor in timing the dispersal phase in pomacentrid species. 20 List of participants Baerwald, Melinda [email protected] Bank, Claudia [email protected] Bensch, Staffan [email protected] Brown, Clare [email protected] Burri, Reto reto.burri@uni-­‐jena.de Caetano-­‐Anolles, Derek [email protected] Chen, Nancy [email protected] Christodoulaki, Eirini [email protected] Curk, Teja [email protected] Delmore, Kira [email protected] Durieux, Gillian [email protected] Ferrer Obiol, Joan [email protected] Frias-­‐Soler, Roberto [email protected]­‐heidelberg.de Gahr, Christoph [email protected] Goldberg, Amy [email protected] Haubold, Bernhard [email protected] Helm, Barbara [email protected] Henrich, Tina [email protected] Jax, Elinor [email protected] Jeschke, Alina [email protected] Jones, Christopher [email protected] Justen, Hannah [email protected] Kaiser, Tobias [email protected] Kishkinev, Dmitry [email protected] Kopp, Gisela [email protected] Kraus, Robert robert.kraus@uni-­‐konstanz.de Lemopoulos, Alexandre [email protected] Liedvogel, Miriam [email protected] Lugo Ramos, Juan S. [email protected] Lundberg, Max [email protected] Marasco, Valeria [email protected] Meek, Mariah [email protected] Menchaca, Angelica [email protected] Merlin, Christine [email protected] Meyer, Britta [email protected] 21 Mueller, Ralf [email protected] Nichols, Krista [email protected] Parody-­‐Merino, Angela [email protected] Partecke, Jesko [email protected] Pérez-­‐Rodríguez, Antón [email protected] Pérez-­‐Tris, Javier [email protected] Pulido, Francisco [email protected] Ranke, Peter S. [email protected] Ringbauer, Harald [email protected] Schalm, Gregor gregor.schalm@uni-­‐oldenburg.de Sekar, Vaishnovi [email protected] Theodosiou, Loukas [email protected] Tietze, Thomas tietze@uni-­‐heidelberg.de Valli, Davide [email protected] van den Hurk, Petrus [email protected] van der Jeugd, Henk [email protected] Van Doren, Benjamin [email protected] van Heusden Jasper [email protected] van Noordwijk, Arie J. [email protected] Voelkl, Bernhard [email protected] von Rönn, Jan [email protected] Wolf, Jochen [email protected] Wotton, Karl [email protected] Zhang, Guojie [email protected] 22 General Information institute
Hotel
train station
Travel You can reach Plön from Hamburg airport by public transport via Kiel or via Lübeck. We recommend to check out your individual travel plan on the official website of B http://www.bahn.de/p_en/view/index.shtml. via Kiel: From 07:15, an hourly shuttle-­‐bus departs from Hamburg airport to Kiel. Tickets can be bought directly in the bus (see also http://www.bahn.de/autokraft/view/angebot/kielius_en/kielius-­‐
ueb.shtml). From Kiel train station you can take the train to Lübeck or Lüneburg, which also stops in Plön. The journey from Hamburg airport to Plön via Kiel takes a little more than 2 hours. via Lübeck: From Hamburg airport on Platform 2 take the Train S1 to Hamburg central station. In Hamburg Airport you can also find ticket machines from DB, where you can buy your ticket for the whole trip to Pl ̈on. From Hamburg central station you then take the train to Lübeck, which usually departs from platform 6. In Lu ̈beck you have to change into the train to Kiel, which also stops in Plön. The journey via Lübeck takes between 2 and 3 hours. 23 Accommodation The Nordic Hotel is located in Öhlmühlenallee 3, 24306 Plön. You can reach it from the train station either by foot or by bus number 331 (stops Danziger Straße or Fegetasche). For further information about the hotel, you can check their website http://www.nordic-­‐hotels.com/en/hotels/ploen-­‐nordic-­‐
hotel-­‐ploen.de Catering Coffee: Between the sessions there will be a coffee break with coffee and snacks. Lunch will be provided in the Cafeteria area of the institute. Dinner: We will have an icebreaker reception and a buffet dinner at the institute on Tuesday. Wednesday and Thursday there will be dinner at a restaurant for the whole group. If you have any questions, ask Kira, Miriam or Chris for help. Lübeck Lübeck is the second largest city of Schleswig-­‐Holstein and is beautifully located at the river Trave. In the middle ages it was an important trading center and leading city of the Hanse, which controlled large shares of the northern European trade. Lu ̈beck has a very nice old town (listed as a world heritage site), the well-­‐known Holstentor and you can also visit the recently built Hansemuseum. • Trains to Lübeck leave every 30 minutes, at hh:15 and hh:45. • Trains back to Plön depart at hh:06 and hh:27 until 21:06 and afterwards at hh:01. The last train leaves Lübeck at 00:17 • The train ride takes about 40 minutes. Kiel Kiel is the capital and most populous city in the northern German state of Schleswig-­‐Holstein. Kiel lies approximately 90 km north of Hamburg. Due to its geographic location in the north of Germany, the southeast of the Jutland peninsula, and the southwestern shore of the Baltic Sea, Kiel has become one of the major maritime centres of Germany. For instance, the city is known for a variety of international sailing events, including the annual Kiel Week, which is the biggest sailing event in the world. The Olympic sailing competitions of the 1936 and the 1972 Summer Olympics were held in Kiel. •
Trains leave every 30 minutes, hh:14 and hh:45. •
Trains back to Plön leave at hh:04 and hh:44. The last train departs in Kiel at 21:43. •
Train ride takes about 31 minutes. WiFi WiFi is accessible for “eduroam” users in the institute. In case you do not have access to ”eduroam”, we will provide you with an account for our guest net, just get in touch with Kira, Miriam or Chris. 24