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Annual Report 2012 Metapopulation Research Group Department of Biosciences Faculty of Biological and Environmental Sciences University of Helsinki Helsinki 2012

Annual Report 2012 - helsinki.fi · Panu Somervuo Ulisses Camargo Mar Cabeza Patrik Koskinen 1 Jussi Jousimo Anni Arponen Anniina Mattila Markku Karhunen 4 Astrid van Teeffelen Swee

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Page 1: Annual Report 2012 - helsinki.fi · Panu Somervuo Ulisses Camargo Mar Cabeza Patrik Koskinen 1 Jussi Jousimo Anni Arponen Anniina Mattila Markku Karhunen 4 Astrid van Teeffelen Swee

Annual Report 2012Metapopulation Research Group

Department of BiosciencesFaculty of Biological and Environmental Sciences

University of Helsinki

Helsinki 2012

Page 2: Annual Report 2012 - helsinki.fi · Panu Somervuo Ulisses Camargo Mar Cabeza Patrik Koskinen 1 Jussi Jousimo Anni Arponen Anniina Mattila Markku Karhunen 4 Astrid van Teeffelen Swee

Contact information

Address: Metapopulation Research Group Department of Biosciences P.O.Box 65 (Viikinkaari 1) FI-00014 University of Helsinki Finland

Phone: +358 9 1911 (exchange)Fax: +358 9 191 57694

E-mail: [email protected]: www.helsinki.fi/science/metapop

© Metapopulation Research Group

Layout: Sami OjanenMRG-logo: Gergely VárkonyiFace photos: Evgeniy Meyke / Sami Ojanen

Printed in Picaset Oy

Helsinki, January 2013

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Contents

& Foreword: Happy researchers .......................................................................5

& Brief history and overview of MRG ...........................................................6

& Research Projects ..................................................................................................9

• The Glanville fritillary model system ................................................................... 10Metapopulation biology of the Glanville fritillary butterfly: Ecological and evolutionary spatial dynamics 10Genomics and genetics of the Glanville fritillary butterfly 12Environmental stress and its effects on life history evolution in wild populations 16

• Metacommunity dynamics ................................................................................... 18Metacommunity dynamics of wood-decaying fungi 18Parasitoid population ecology 20

• Coevolutionary dynamics and radiations ........................................................... 22Species Interactions in Metapopulations 22 Evolutionary radiations of dung beetles in Madagascar 24

• Mathematical ecology ............................................................................................ 26Modelling dispersal and population dynamics 26European Boreal Forest Biodiversity (EBFB) 28

• Systematic conservation planning ....................................................................... 30Biodiversity conservation informatics 30Conservation Effectiveness 34Climate change 36

& Supporting personnel ...................................................................................... 38

& Synopsis of the year 2012 ................................................................................ 41

Publications 42Honours and awards 49Visitors 49Teaching 50Annual meeting in St Petersburg 52Budget 53

& Prospects for the year 2013 .......................................................................... 52

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Research groups are unusual work places, though I have to admit that I say so without knowing what any other work place would really be like. I believe that research groups are less hierarchical than most other work places, and very democratic in the sense that great ideas are always appreciated regardless of who puts them forward. Research groups tend to consist of individuals from novices (undergraduate students) to real experts, and many research groups, like our MRG, consist of students, researchers and supporting personnel with widely different backgrounds and nationalities. Most importantly, research groups are unusual work places because research is such an unusual occupation. We are in the business of making discoveries, pushing the frontiers of knowledge further and further. The challenge is to tolerate the uncertainty and the inevitable failures that are part of the exploration; the reward is the satisfaction that accompanies success, whether it is the PhD thesis completed, a new research paper, or the solution to something that you spent a lot of time in figuring out. However, as a starting researcher, you should not expect an easy ride from one success to another. If you do, the chances are that you will find yourself, sooner or later, doing something else in your life. Young scientists have to learn to get sufficient satisfaction from small enough accomplishments. Young scientists have to learn to love science, and especially their own science. If you don’t believe that what you are doing is really, really exciting and important, why should the others think so?

Sometimes a researcher stumbles to results that were unexpected and which appear to offer a glimpse of a new horizon for research. I had this feeling earlier this year while examining possible links between environmental biodiversity around people’s homes, the composition of the bacterial community (microbiota) on their skin, and their atopic condition (level of specific IgE anti bodies; Hanski et al. 2012, Proc Natl Acad Sci US 109, 8334-8339). It was a pleasure to work with enthusiastic researchers with very different backgrounds, from ecology to immunology, from allergy studies to metagenomics and bioinformatics, all passionately excited about what we are doing. Science is cool!

Research groups are unusual work places because of the mobility of especially the younger members. Many new students and researchers have joined MRG in 2012. Our two new students are Ulisses Camargo from Brazil and Wolfgang Reschka from Austria. The new post docs include Enrico Di Minin from Italy, via UK; Kristina Karlsson Green from Sweden; Guillaume Blanchet from Canada; Benoit Barres from France; and Anton Chernenko from Russia (PhD in Helsinki). Others have left us, moved on in their life and career: Chris Wheat moved to Sweden; Emily Hornett and Maaike de Jong to UK; Charlotte Tollenaere to France and Tommi Mononen moved to Aalto University in Espoo. Very exceptionally, we had only one PhD student finishing this year, Heini Kujala, who left to start a post doc in Australia.

So research groups are special work places – and MRG is a particularly special, and happy, research group, because we have such a great team of supporting personnel: Viia, Sami, Krista, Johanna and Aija in the office, Toshka, Annukka, Heini, Alison and Pia in the molecular laboratory, Suvi in charge of the butterfly laboratory at the Lammi biological station, and Evgeniy developing data management with EarthCape. Viia and Sami in our office are the key players in our team, without whom this research group would not be the happy community that we are.

Ilkka Hanski

Happy researchers

Foreword

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MRG -ANNUAL REPORT 20126 MRG -ANNUAL REPORT 20126

Brief history and overview of MRG

Metapopulation Research Group was established by Ilkka Hanski 21 years ago, in 1991. Ilkka had worked on the ecology of

spatially structured populations since the late 1970’s. The early work was concerned with small-scale spatial aggregation of individuals within populations and how that might affect the coexistence of competitors. Since the early 1980’s the focus had shifted to larger spatial scales and to the dynamics of metapopulations, networks of local populations with relatively independent demographic dynamics. Significant events leading to the establishment of MRG included the first international workshop on metapopulation ecology organized by Ilkka and Michael Gilpin (San Diego, US) in 1989, which resulted in the first edited volume on the subject (Gilpin & Hanski, 1991, Metapopulation Dynamics: Empirical and Theoretical Investigations, Academic Press, London). The long-term metapopulation study of the Glanville fritillary butterfly in the Åland Islands in Finland was started in 1991.

Current research

MRG is the leading research group worldwide in metapopulation biology and one of the Centres-of-Excellence in research nominated by the Academy of Finland (national research council) for 2000-2005, 2006-11 and again for 2012-2016. Our past strengths include successful integration of theory, modeling and empirical studies in the same projects. The Glanville fritillary butterfly has become a widely recognized model system in population biology, but other long-term projects are increasing in prominence: the host-parasitoid system studied by Saskya van Nouhuys, the

plant-mildew system studied by Anna-Liisa Laine, the community of wood-decomposing fungi studied by Otso Ovaskainen, and the radiation of endemic dung beetles in Madagascar studied by Ilkka. Atte Moilanen’s Zonation software has attained worldwide reputation in systematic conservation planning. In the past, MRG was primarily a research group in ecology, but this is no longer the case. The senior researchers have opened up new fronts of research to bring together population biology and state-of-the-art mathematical modeling (Otso Ovaskainen), molecular biology (Mikko Frilander) and genomics (Rainer Lehtonen), as well as developed new approaches to conservation biology and reserve design (Atte Moilanen, Mar Cabeza). The metapopulation concept is no longer as fundamental to our research than it was in the past, though processes related to the spatial structure and dynamics of populations remain the focus of much of our research.

Current structure

The graph shows the growth of MRG since 1992. At present, MRG is a highly international group of 73 researchers (9), post docs (28), post graduate students (23), and supporting personnel (13) representing 19 different nationalities. MRG currently consists of seven research groups with their own students and post docs and largely own funding. We are united by the shared Centre-of-Excellence funding, which supports a common office and other facilities. Equally, we are united by much interaction among the research groups around four major research themes:

I Local adaptation, ecological and coevolutionary dynamics, and evolutionary radiations

II Genomics, genetics and functional molecular biology

III Mathematical ecologyIV Ecological decision analysis and applied

conservationThe table on the next page lists the group leaders and senior researchers (in bold), independent researchers and post docs (in italics) and post graduate students.

Theme I – Local adaptation, ecological and coevolutionary dynamics, and evolutionary radiations The objective is to advance the general understanding of the dynamics of species living in heterogeneous environments, with explicit attention to interactions

Fig. 1. MRG personnel since 1992.

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Overview

7

between demographic dynamics and local adaptation, between behaviour and ecological dynamics in metacommunities, and between the dynamics of species’ geographical ranges and evolutionary radiations. Many projects involve molecular studies and modelling.

Theme II – Genomics, genetics and functional molecular biologyThe main objective is to develop and implement genomic and genetic tools to the study of natural populations of non-model species. Most of the work is focused on the Glanville fritillary, but other study projects also benefit of the methods developed. The current main task is to complete the draft genome of the Glanville fritillary, which will greatly facilitate research on this major study system in the future.

Theme III – Mathematical ecologyThis theme has four objectives. First, we aim to develop general theory in spatial ecology, genetics and evolution.

Second, we aim to relate these theories to empirical data via new statistical approaches. Third, many of the empirical projects described in Themes I and II will be complemented with more specific modelling projects. Fourth, we will develop modelling tools to better connect decision analysis and conservation (Theme IV) to basic population and community ecology.

Theme IV – Ecological decision analysis and applied conservationThis theme has four objectives, all building on past research by the senior researchers in MRG. First, we develop methods for conservation prioritization that integrate across multiple environments and levels of biological organization. Second, we consolidate the linkage between conservation planning and cutting-edge species-level and community-level spatial modelling. Third, we improve the understanding of how climate change should be accounted for in conservation. Fourth, we develop methods for the evaluation of conservation outcomes.

Ilkka Hanski Otso Ovaskainen Atte MoilanenRainer Lehtonen Guillaume Blanchet Enrico Di MininVirpi Ahola Henjo De Knegt 3 Jussi LaitilaAnton Chernenko Maria Delgado Federico PouzolsMaaike De Jong Juri Kurhinen Tuuli ToivonenAnne Duplouy Juho Pennanen Peter KullbergAndreia Miraldo Dmitry Schigel Joona Lehtomäki 3

Pasi Rastas Daniel SimpsonPanu Somervuo Ulisses Camargo Mar CabezaPatrik Koskinen 1 Jussi Jousimo Anni ArponenAnniina Mattila Markku Karhunen 4 Astrid van TeeffelenSwee Chong Wong Sonja Koskela Silvija Budaviciute

Veera Norros Johanna EklundAnna-Liisa Laine Tanjona Ramiadantsoa 3 Henna FabritiusBenoit Barres Rachel GarciaKristina Karlsson 2 Saskya van Nouhuys Heini KujalaHannu Mäkinen Delia Pinto Zevallos Laura MellerAyco Tack Christelle Couchoux Maria Triviño 5

Charlotte Tollenaere Wolfgang Reschka Laure ZupanRiikka AlanenHanna Susi Mikko Frilander Chris Wheat

Jouni Kvist 3 Emily HornettMarjo Saastamoinen + 4 post docs/PhD students in Institute of Biotechnology

1 with Liisa Holm, 2 with Saskya van Nouhuys, 3 in collaboration with Ilkka Hanski, 4 with Juha Merilä, 5 with M.B. Araújo,

Table 1. Subgroup leaders / senior researchers (bold), researchers/post docs (italics) and PhD students in MRG.

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Research Projects

The Glanville fritillary model system

Metapopulation biology of the Glanville fritillary butterfly: Ecological and evolutionary spatial dynamics

Genomics and genetics of the Glanville fritillary butterflyEnvironmental stress and its effects on life history evolution in

wild populations

Metacommunity dynamics

Metacommunity dynamics of wood-decaying fungiParasitoid population ecology

Coevolutionary dynamics and radiations

Species Interactions in MetapopulationsEvolutionary radiations of dung beetles in Madagascar

Mathematical ecology

Modelling dispersal and population dynamicsEuropean Boreal Forest Biodiversity (EBFB)

Systematic conservation planning

Biodiversity conservation informaticsConservation EffectivenessClimate Change

Supporting personnel

© Sami Ojanen

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10 MRG -ANNUAL REPORT 2012

Ilkka HanskiProject leader

Metapopulation biology of the Glanville fritillary butterfly: Ecological and evolutionary spatial dynamics

Anniina MattilaPhD -student

The large metapopulation of the Glanville fritillary butterfly (Melitaea cinxia) in the Åland Islands in southwestern Finland has

been studied since 1991. Over the years, the Glanville fritillary has become a widely recognized model system not only in metapopulation ecology but in population biology more generally. The research has made many contributions to the study of the ecological, genetic and evolutionary consequences of habitat fragmentation, and the empirical studies have stimulated the development of new concepts and models. Currently the ecological studies are coupled with functional genomics research (p. 12).

Fig. 1. Genetic load and heterosis in a small isolated population (PT) of the Glanville fritillary butterfly. A. Egg hatching rate (%) in PT (gray) and ÅL (white, large reference population) and in crosses between PT females and males from other regional populations (hatched gray lines). B. Peak flight metabolic rate (residual from a regression against adult weight) in field-collected PT (gray) and ÅL (white) females, and in the female offspring of crosses between PT females and males from other regional populations (gray lines). These results show instant and complete fitness recovery in the crosses (Mattila et al.2012).

Maaike de JongPost docAnne Duplouy

Post doc

Highlights of the year

Anne Duplouy has conducted a common garden study of the life history ecology and evolution of four regional populations of the Glanville fritillary from two highly fragmented landscapes (Åland Islands in Finland and Uppland in Sweden) and two continuous landscapes (Saaremaa Island in Estonia and Öland Island in Sweden). As predicted by previous models, butterflies from the fragmented landscapes have higher dispersal rate (as measured by peak flight metabolic rate) than butterflies from continuous landscapes. Anniina Mattila and others have done research on a completely isolated,

PTxPT Cross ÅLxÅLPopulation/cross

PT Cross ÅLPopulation/cross

BA

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The Glanville fritillary model system

Collaborators

Prof Rongjiang Wang, Peking University, ChinaProf Paul Brakefield, University of Cambridge, UKDr Kristian Niitepõld, Stanford University, USAProf Patrik Nosil, University of Sheffield, UK

Selected publications

Hanski I. (2012). Metapopulations and spatial population processes. In Oxford Bibliographies in Ecology. David Gibson (ed.). New York: Oxford University Press. Entry Launch May 2012.

Hanski I. (2012). Eco-evolutionary dynamics in a changing world. In The Year in Ecology and Conservation Biology. Eds. R.S. Ostfeld and W.H. Schlesinger. Annals of the New York Academy of Sciences 1249, 1-17.

Niitepõld K. and Hanski I. (2013). A long life in the fast lane: positive association between peak metabolic rate and lifespan in a butterfly. Journal of Experimental Biology, doi:10.1242/jeb.080739.

Mattila A.L.K., Duplouy A., Kirjokangas M., Lehtonen R., Rastas P. and Hansk, I. (2012). High genetic load in an isolated butterfly population. Proc. Natl. Acad. Sci. US, 37, E2496-E2505.

Hanski I. (2012). Dispersal and eco-evolutionary dynamics in the Glanville fritillary butterfly. In Dispersal Ecology and Evolution Clobert J., Baguette M., Benton T.G. and Bullock J.M. (eds.), pp. 290-303. Oxford University Press, Oxford.

Fig. 2. A) Distance (cm) to an air source, and the corresponding force of the air flow, at the point when the butterfly lost its grip, for young (<5 days) and old butterflies from Åland (gray boxes) and PT (empty boxes). B) The angle of curvature of the claw in butterflies from Åland and in PT.

Fig. 3. Cyp allele frequency is significantly correlated with average host plant preference in the Åland Islands (average values for different parts of the study area). Here, the horizontal axis shows the frequency of the G allele in Cyp and the vertical axis shows preference for Plantago over Veronica.

small and old (at least 76 years) local population of the Glanville fritillary on the island of Pikku Tytärsaari (PT) in the Gulf of Finland. Despite its high genetic load manifested as greatly reduced fitness (Fig. 1), one striking local adaptation has been documented for the PT population. The claws of the PT butterflies have a greater curvature, apparently improving their grip and allowing the butterflies to resist strong air force, and hence windy conditions on the island, which is expected to reduce the risk of being blown off to the sea (Fig. 2). Maaike de Jong has studied heritability of life-history traits and fitness components in an experiment lasting for two generations. This study revealed a high heritability for lifetime reproductive success, the key component of fitness. A large part of this heritability was explained by a single Cytochrome P450 gene (Cyp), which is known to be involved in host plant adaptation in other insect species. The results show a strong

correlation between Cyp and host plant preference across the Åland Islands (Fig. 3). It may well turn out that the dynamics of genetic variation in the Cyp gene is coupled with ecological dynamics and hence represents yet another example of eco-evolutionary dynamics in the Glanville fritillary. The models of eco-evolutionary dynamics that have been stimulated by the Glanville fritillary project have been recently applied to another study system, the Timema walking stick insects in California, in collaboration with Prof Patrik Nosil.

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Rainer LehtonenProject leader

Genomics and genetics of the Glanville fritillary butterfly

The main objective of this research project is to develop and implement genomic and genetic tools for the study of natural populations of

non-model species. Most of the work is focused on the Glanville fritillary, but other study projects will benefit of the methods to be developed. We aim at localizing and identifying genes and genetic variants affecting phenotypic and life-history traits that influence local adaptation and population dynamics. A population-wide pedigree will be constructed for the Glanville fritillary using spatial and genetic data.

Highlights of the year

1. We have completed the sequencing of the ~400 Mb genome of the Glanville fritillary using Roche 454, Illumina GAII/HiSeq2000 and SOLiD4/5500

Ilkka HanskiSenior researcher

Virpi AholaPost doc

Mikko FrilanderSenior researcher

Pasi RastasPost doc

Supporting personnel: Toshka Nyman, Alison Ollikainen, Marja-Leena Peltonen, Annukka Ruokolainen, Suvi Saarnio, Pia Välitalo

Patrik KoskinenPhD -student

Panu SomervuoPost doc

Jouni KvistPhD -student

Swee Chong WongPhD -student

platforms. The current draft genome assembly (table 1, figure 1) consists of 8,262 scaffolds (>1,500 bp) and includes approximately 15,000 gene models. Automated annotations have been manually curated for more than 600 genes. Roche-Nimblegen and SOLiD RAD-tag data from controlled crosses including 22,000 SNPs have been used to construct a genetic map with the correct number of chromosomes (n=31). The 1st version of the genome will be published in 2013.

2. RNA sequencing of population samples and samples from experiments on flight metabolic rate. We have individually sequenced transcriptomes (RNA) of 198 samples originating from five different regional populations. Åland and Uppland populations live in a fragmented landscape, whereas Saaremaa and Öland represent

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The Glanville fritillary model system

Table. 1. Assembly and validation statistics for the Glanville fritillary genome. v1 and v1.1 refer to the two different versions of the assembly. N50 is the median length of contigs/ scaffolds in the assembly. CEGMA (Core Eukaryotic Genes Mapping Approach) dataset consists of 248 eukaryotic proteins that are conserved over most of the taxa. “Complete” refers to the Glanville fritillary predicted proteins which give an alignment length minimum of 70% to the reference protein cluster. In the validation based on the genetic map, SNPs within a scaffold (or contig) are counted to be correct if they map to one chromosome but are assumed to involve assembly errors if mapped to two or more different chromosomes.

continuous landscapes. Pikku Tytärsaari is a small isolated population suffering from a high genetic load. An experiment on flight metabolism and differences in flight performance includes 90 RNA samples. In-depth analyses of these data sets, including Gene Ontology (figure 2) and pathway enrichment analyses, are on-going.

3. The genomic information has been used to design genotyping assays for large-scale association and linkage studies. Targeted genotyping experiments have revealed many novel gene-phenotype associations such as a vitellin-degrading protease precurson gene in larval growth (Ahola et al. in prep.) and the occurrence of an extra larval instar (Saastamoinen et al. in press), and a cytochrome P450 gene affecting the reproductive performance of Glanville fritillary females (de Jong et al. in prep.).

Fig. 1. Influence of library insert size on scaffold N50 size. The most significant increase in scaffold size occurred after the 454 20kb library data (library 9) had been added to the assembly.

CollaboratorsPetri Auvinen and Lars Paulin, Institute of Biotechnology,

University of Helsinki (sequencing and assembly)Liisa Holm, University of Helsinki (annotation)Minna Taipale, Karolinska Institute, Sweden (Illumina

sequencing)Daniel Lawson, EBI, UK (annotation of the full genome)Veli Mäkinen and Esko Ukkonen and their groups,

University of Helsinki (new solutions for de novo genome assembly)

Markus Perola and Samuli Ripatti, National Institute of Health and Welfare, Finland (association analyses of complex traits)

Päivi Lahermo, Institute of Molecular Medicine Finland (genotyping).

IT Center for Science (CSC), Finland (supercomputing resources)

Publications

Mattila A.L.K., Duplouy A, Kirjokangas M., Lehtonen R., Rastas P. and Hanski I. (2012). High genetic load in an old isolated butterfly population. Proc. Natl. Acad. Sci. 109(37), E2496-505.

Kvist J., Wheat C.W., Kallioniemi E., Saastamoinen M., Hanski I. and Frilander M.J. (2013). Temperature treatments during larval development reveal extensive heritable and plastic variation in gene expression and life history traits. Mol Ecol. 22:602-619

Saastamoinen M., Ikonen S., Wong S.C., Lehtonen R. and Hanski I. Plastic larval development in a butterfly has complex environmental and genetic causes and consequences for population dynamics. J. Anim. Ecol., in press.

Karinen S., Saarinen S., Lehtonen R., Rastas P., Vahteristo P., Aaltonen L.A. and Hautaniemi S. 2012. Rule-based induction method for haplotype comparison and identification of candidate disease loci. Genome Med. 4(21).

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14 MRG -ANNUAL REPORT 2012

Fig. 2. Gene Ontology (GO) enrichment analysis of flight-induced genes. The experiment included male and female butterflies originating from Åland and Pikku Tytärsaari populations. Thorax tissue was collected 1 or approximately 10 hours after flight (n=61) and from control individuals (n=29) for RNA sequencing. Out of 8221 genes expressed in the experiment, differential expression of 755 genes was attributed to the flight treatment with false discovery rate 0.05. GO annotation was available for 441 genes, which were used in the enrichment analysis. Hierarchical tree for the significantly enriched (P value < 0.05) GO groups are shown. Biologically promising GO groups are highlighted, including for example hypoxia, carboxylic acid metabolism, translation and rRNA processing. The information in the nodes is: GO group ID, GO group description, enrichment P value and the number of Glanville fritillary genes mapped to the GO group/total number of genes in the group.

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The Glanville fritillary model system

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Environmental stress and its effects on life history evolution in wild populations

The ability to cope with, react or even adapt to environmental stress is essential for most organisms. Yet, surprisingly little is still known

about the strategies and genetic mechanisms involved in coping with environmental stress in wild populations. In this project, I have studied the influence of stressful developmental conditions, in terms of thermal and resource levels, on life history variation in two butterfly species, Bicyclus anynana and Melitaea cinxia. With the help of quantitative genetics and candidate gene approaches I have further aimed to understand the (genetic) mechanisms underlying the observed responses.

Marjo SaastamoinenResearcher

Highlights of the year

The 2012 was the final year of my Post-doctoral research grant project funded by the Academy of Finland. With this project I have shown how the two study species B. anynana and M. cinxia respond to developmental nutritional stress in a very different way. Interestingly, however, in both species individuals seem to use developmental conditions as predictive cues of the conditions they or their offspring are likely to encounter later on in life (i.e. predictive adaptive response). In B. anynana the response to developmental stress is a change in resource allocation patterns between thorax and abdomen, which results in increased flight ability of the stressed individuals. This response potentially allows the stressed individuals to move away from the poor habitat patches, as was shown by our theoretical model1. In M. cinxia, on the other hand, mothers stressed during development produce offspring that can cope better with similar type of stress during their own development compared with offspring from non-stressed mothers2 (Fig. 1). To compensate for suboptimal thermal conditions during development, M. cinxia can even increase the number of developmental instars3. Variation in instar number seems to be

additionally associated with variation in three candidate genes. In general, it seems evident that genotype*environment interactions are very important determinants of life history variation in both of these species.

In 2012, I also initiated research on assessing the link between dispersal and immune defence in the Glanville fritillary butterfly (Fig. 2.). In May I had a wonderful experience to conduct a pilot experiment in collaboration with Jean Clobert and Michel Baguette in the experimental-metapopulation, the Metatron (Fig. 3), in Moulis, France.

Fig. 1. Direct and trans-generational effects (± s.e.) on pupal mass. Current developmental treatment is represented by the x-axes and maternal treatment by the different lines. (Modified from Saastamoinen et al., Oecologia).

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The Glanville fritillary model system

Collaborators

Michelle Baguette, Experimental Ecology Research Centre, CNRS in Moulis, France

Prof. Paul Brakefield, University of Cambridge, UK Jean Clobert, Experimental Ecology Research Centre,

CNRS in Moulis, France Jon Brommer, University of TurkuMarkus Rantala, University of Turku Prof. Bas Zwaan, University of Wageningen, The

Netherlands

Fig. 3. Experimental metapopualtion (CNRS in Moulis, France).

Selected publications

1. van den Heuvel J., Saastamoinen M., et al. (2013). The predictive adaptive response: modeling the life history of a tropical butterfly (Bicyclus anynana). American Naturalist, in press.

2. Saastamoinen M., Norio H. & Van Nouhuys S. (2013). Direct and trans-generational responses to food deprivation during development in the Glanville fritillary butterfly. Oecologia, in press.

3. Saastamoinen M., Ikonen S., Wong S.W., Lehtonen R. & Hanski I. (2013). Plastic larval development in a butterfly has complex environmental and genetic causes and consequences for population dynamics. Journal of Animal Ecology, in press.

Fig. 2. Assessment of encapsulation rate in an adult M. cinxia.

©Linda Peltola

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18 MRG -ANNUAL REPORT 2012

Otso OvaskainenProject leader

Metacommunity dynamics ofwood-decaying fungi

Sonja KoskelaPhD -student

Veera NorrosPhD -student

This project is focused on metacommunity dynamics of wood-inhabiting fungi living

in the dynamic habitat provided by decaying trees. Wood-inhabiting aphyllophoroid fungi (Basidiomycota) are a diverse, ecologically important and in Fennoscandia taxonomically well known group of forest species. Many species in this group are red-listed, mainly due to large-scale reduction in the area of natural forests and the amount of dead wood in all forests. We and others have shown that in fragmented landscapes some dead-wood dependent fungi have declined more than what would be expected from the loss of their microhabitats. Such a non-linear response indicates the presence of regional extinction thresholds, but the exact mechanisms behind species declines are still poorly understood. Our aim is to find out what drives the dynamics of this species community, and consequently what kind of management measures would be most effective in improving the viability of wood-inhabiting fungi in boreal forests. One key aspect of the work is molecular identification of fungi from next generation sequencing data, and the combination of sequencing data with sporocarp survey data.

Technician: Heini Ali-Kovero

Fig. 1. Woodpecker´s job. To avoid felling trees in protected areas, sampling standing trees requires ropes and carabiners (left). Samples from living trees (above) often contain a mix of hard sawdust (light colour) and decayed particles (brown), suggesting presence of wood-decaying fungi.

Dmitry SchigelPost doc

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Metacommunity dynamics

Collaborators

Dr. Üllar Rannik and Dr. Tareq Hussein, University of Helsinki

Dr. Panu Halme, University of JyväskyläDr. Raisa Mäkipää, Dr. Reijo Penttilä, and M.Sc. Juha

Siitonen, Finnish Forest Research InstituteProf. Jan Stenlid, M.Sc. Elisabet Eriksson, the Swedish

University of Agricultural SciencesDr. Jenni Nordén and Dr. Karl-Henrik Larsson, University

of OsloDr. Henrik Nilsson, University of Gothenburg

Recent publications

Liu X., Suzuki A. And Schigel D. (2012) Editorial: The impact of fungi on other organisms. — Mycology 3: 1.

Nilsson R. H., Ovaskainen O., et al. (2012). Five simple guidelines for establishing basic authenticity and reliability of newly generated fungal ITS sequences. MycoKeys 4, 37-63.

Norros V., Penttilä R., Suominen M. and Ovaskainen O. (2012). Dispersal may limit the occurrence of specialist wood decay fungi already at small spatial scales. Oikos 121, 961-974.

Schigel D.S. (2012). Fungivory of saproxylic Coleoptera: the mystery of rejected polypores. In: Saproxylic beetles in Europe: monitoring, biology and conservation. — Studia forestalia Slovenica 137: 53–58.

Schigel D.S. (2012) Fungivory and host associations of Coleoptera: a review of research approaches. — Mycology, 3:258-272.

Fig. 2. Laboratory measurements on fungal spore deposition. Panel (a) shows the experimental setup which Veera used to study the deposition of fungal spores into canopy structures. The spores produced by the fruit body (1) went through a pipe (7) stuffed with spruce needles. The penetration rate was estimated by comparing spore concentrations measured by optical particle counter (9) before and after the obstacle. Panel b shows how the measured penetration rate decreases with increasing flow speed and increasing needle density for two fungal species with different spore sizes.

Highlights of the year

Dmitry obtained a Finnish Academy post doc grant for the project ”Colonization gates and establishment of wood-decaying fungi in European Spruce”. This year he sampled living and standing dead trees (Fig 1) to reconstruct the history of fungal arrival by spores. In addition to field work in Finland, Dmitry collected samples from China, Poland, and Sweden. Supervised by Dmitry, Virve Koivuranta started her master´s project on insects as dispersal vectors of wood-decaying fungi.

At the time of writing this report, Veera is busy with finishing her PhD, the deadline for submitting the work to pre-examiners being early January 2013.The combination of field measurements, laboratory experiments and mathematical modelling has resulted in many interesting results. In particular, we have been long interested to learn about the role of spore size in fungal dispersal. The collaboration with aerosol physicists has shown that deposition processes (rather than e.g. gravitational settling) seems to be the key in making large spores disperse shorter distances than small spores (Fig. 2).

Sonja continued to develop probabilistic methods of taxonomic classification necessary for species-level identification based on molecular barcoding data - such data are now being generated in large quantities thanks to Heini’s work.

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20 MRG -ANNUAL REPORT 2012

PhD -student: Katie Sullivan (Cornell University)Undergraduate students: Aapo Ahola1, Riikka Alanen1, Peter Kullberg1, Terhi Lahtinen1, David Muru, Markus Tietäväinen1 co-supervised with Anna-Liisa Laine

Saskya van NouhuysProject leader

Parasitoid population ecology

We study the community, population and behavioural ecology of parasitoids in a spatial context. The project started in the

early 90s with the spatial dynamics of two parasitoids of the Glanville fritillary butterfly in Åland, Finland. Gradually it has broadened to include other parasitoids, hyperparasitoids, pathogens, symbionts, the host food plants, and a related butterfly, Melitaea athalia.

Highlights of the year

A highlight from the population ecology side of our research comes from a study of parasitoid virulence and host susceptibility across five Baltic populations of the Glanville fritillary butterfly. The ability to resist parasitism differed among butterfly populations, and

Fig. 1. The proportion of C. melitaearum in each immature life stage at host diapause stage, separated by parasitoid origin (a) and host origin (b). In each pie, the fraction maturing in the autumn (two dark pie slices: cocoons and third instar larvae) is offset from the fraction maturing in the spring (two light pie slices: first and second instar larvae). * indicates a significant difference at p < 0.05 in the fraction maturing in autumn.

Christelle CouchouxPhD -student

aspects of parasitoid (Cotesia melitaearum) virulence, such as brood size, differed among parasitoid populations. There was however, no evidence of local adaptation by either species. There was also no apparent cost of inbreeding for the parasitoid from the tiny isolated island of Pikku Tytärsaari. One interesting trait that differed among populations for both hosts and parasitoids was the number of wasp generations per year (Fig. 1), which can have a large impact on their population dynamics.

Previously we have studied the multitrophic effects of toxic chemicals produced by plants on parasitoids. This year we started working on another aspect of plant chemistry, the volatile organic chemicals (VOCs) that the plants emit. Since herbivores and their parasitoids both use these volatiles as foraging cues, plants are under selection to make them both inconspicuous to herbivores

and attractive to parasitoids. Delia Pinto found that a plant that has been fed upon by M. cinxia has a much different VOC profile than an intact plant, and that even oviposition by M. cinxia onto the surface of a leaf causes the volatile emission to change (Fig. 2). The next steps are

Delia PintoPost doc

Wolfgang ReschkaPhD -student

a) parasitoid origin

UpplandSaaremaaPikku-TytärsaariÅland

b) host origin

Åland UpplandSaaremaaÖland

2nd instar 1st instar

cocoons

3rd instar* *

*

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Metacommunity dynamics

to measure the responses of M. cinxia and the parasitoids to the different volatile profiles.

We have also continued a long-term study of the behavioural ecology of the parasitoid Hyposoter horticola. The adult females of this species compete strongly, with multiple females finding and monitoring host egg clusters before they are ready to be parasitized. A new graduate student, Wolfgang Reschka, is studying how the wasp keeps track of the spatial locations of host egg clusters in a landscape. He will do this by following the foraging routes of individual wasps using RFID chip technology. The first step was to figure out what sized chip can be glued to a wasp, still allowing it to fly and forage (Fig. 3).

After parasitizing a fraction of the hosts in a cluster an individual H. horticola leaves a deterrent mark on the plant. This year Christelle Couchoux used microsatellite markers to determine the effectiveness of the deterrent. She found that while one mother wasp parasitizes the majority of hosts in a cluster there are, as expected in the competitive environment, “cheaters,” especially in the large host clusters (Fig. 4).

Collaborators

Dr. H. Hakola, Finnish Meteorological Institute, FIDr. J. A. Harvey, Netherlands Institute of Ecology, NLProf. J. K. Holopainen, University of Eastern Finland, FIProf. R. F. Medina, Texas A & M University, USA Prof. D. Quicke, Imperial College, Silwood Park, UKDr. J. H. Reudler, University of Jyväskylä, FIDr. M. S. Shaw, National Museum of Scotland, UK

Recent publications

van Nouhuys S., Niemikapee S., Hanski, I. (2012). Variation in a host-parasitoid interaction across independent populations, Insects, special issue on insect Immunity. Insects, 3: 1236-1256

van Nouhuys S. & Kraft T.S. (2012). Indirect interaction between butterflies meditated by a shared pupal parasitoid Hyposoter horticola. Population Ecology, 54: 251-260 .

Saastamoinen M., Hirai N., van Nouhuys S. (2013). Direct and trans-generational responses to food deprivation during development in the Glanville fritillary butterfly. Oecologia,171:93-104.

van Nouhuys S, et al. (2012). Performance of secondary parasitoids on chemically defended and undefended hosts. Basic and Applied Ecology, 13: 241-249

Fig. 3. The parasitoid wasp Hyposoter horticola with a small piece of metal glued to its thorax.

Fig. 2. Emissions rate of 9 volatile compounds from Veronica spicata that is undamaged (white bars), fed upon (gray bars), and oviposited upon (black bars) by Melitaea cinxia. Significance between treatments is shown as * P < 0.05; ** P < 0.05; *** P < 0.001.

Fig. 4. The number of Melitaea cinxia larvae parasitized by Hyposoter horticola (coloured stripes) and unparasitized (gray stripes) in 10 egg clusters parasitized in the field. In each bar the progeny of each mother wasp is represented by a different colour, so for example all of the parasitoid offspring in cluster F have the same mother whereas in cluster I most progeny were mothered by a single wasp, but 7 other mothers produced a few progeny each. The parentage of the offspring is determined using 14 microsatellite markers. The yellow colour in some of the bars represents wasp progeny that could not be genotyped (due to poor quality DNA extraction), and therefore could not be assigned to a mother.

*

**

***

**

**

**

**

Melitaea cinxia egg cluster in order of size

Num

ber o

f cat

erpi

llars

0

200

150

100

50

A B C D E F G H I J

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22 MRG -ANNUAL REPORT 2012

Anna-Liisa LaineProject leader

Species Interactions in Metapopulations

Our work is focused on uncovering eco-evolutionary links in a large natural plant-pathogen metapopulation of

the powdery mildew fungus, Podosphaera plantaginis, infecting Plantago lanceolata in the Åland Islands.

In 2012 our group expanded as first Krista Raveala began work as a research technician in January. Riikka Alanen turned in her MSc thesis in April and has started her PhD work focused on overwintering of Po. plantaginis. During spring term, Benoit Pernechele (Louvain Academy, Belgium) completed his thesis project under the supervision of Charlotte. In May Kristina Karlsson-Green was awarded the Formas grant in Sweden and she joined us to carry out a joint project with Saskya van Nouhuys and Marjo Saastamoinen on the interactions among trophic levels associated with Pl. lanceolata. During a post doctoral project Hannu Mäkinen developed a qPCR method for detecting and quantifying

Fulbright scholar: Felix HornsUndergraduate students: Aapo Ahola1, Peter Kullberg1, Terhi Lahtinen1, Sini Mursinoff 2, Benoit Pernechele3

* Joint project with Marjo Saastamoinen & Saskya van Nouhuys1 Supervisors Anna-Liisa Laine & Saskya van Nouhuys2 Supervisors Anna-Liisa Laine & Ayco Tack3 Supervisor Charlotte Tollenaere

Ayco TackPost doc

Charlotte TollenaerePost doc

Hanna SusiPhD -student

Benoit BarrèsPost doc

Kristina Karlsson Green *Post doc

Hannu MäkinenPost doc

Riikka AlanenPhD -student

0.1

1.0

10.0

0 50 100 150 200 250Distance from source (cm)

Num

ber o

f spo

res

Group sizeTotal12

3−5>5

Fig. 1. The number of spores landing on microscope slides decreases with distance from the pathogen source. The black line shows the total number of spores and the coloured lines show the contribution of different spore group sizes.

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Coevolutionary dynamics and radiations

Collaborators:

Dr Jeremy Burdon, CSIRO Plant Industry, Canberra, Australia

Prof. David Gadoury, University of Cornell, USAEmilie Haon-Lasportes, INRA, Avignon, FranceProf. Levente Kiss, Academy of Sciences, Budapest,

HungaryDr Samuel Soubeyrand, INRA, Avignon, FranceDr Peter Thrall, CSIRO Plant Industry, Canberra, Australia

Key Publications

Tollenaere C., Susi H., Nokso-Koivisto J., Koskinen P. Tack A. J. M., Auvinen P. Paulin L., Frilander M. J., Lehtonen R. & Laine A.-L. (2012). SNP Design from 454 Sequencing of Podosphaera plantaginis Transcriptome Reveals a Genetically Diverse Pathogen Metapopulation with High Levels of Mixed-Genotype Infection. PLoS ONE, 7:e52492.

Tack A., Thrall P. H. Barrett L. G., Burdon J. J. & Laine A.-L. (2012). Variation in infectivity and aggressiveness in space and time in wild host-pathogen systems - causes and consequences. Journal of Evolutionary Biology 25:1918-1936.

Nemri A., Barrett L. G., Laine A.-L., et al. (2012). Mode of reproduction predicts population structure at different spatial scales in the Linum marginale - Melampsora lini association. PLoS ONE 7: e41366.

Thrall P. H., Laine A.-L., et al. (2012). Rapid genetic change underpins antagonistic coevolution in a natural host-pathogen metapopulation. Ecology Letters, 15: 425-35.

co-infection. And finally, Benoit Barrès joined us in October to begin work on molecular epidemiology of Po. plantaginis.

Highlights of the year

• This year marked the highest disease prevalence in the Åland metapopulation with nearly 700 infected populations.

• The project on Po. plantaginis transcriptome sequencing, and development and validation of the first SNP panel was completed this year through a successful collaboration of many partners (Tollenaere et al. 2012). We discovered that approximately half of the local pathogen populations contain genetic diversity, and in these populations co-infection is common.

• Ayco’s dispersal experiments using a wind tunnel and common garden setting revealed that each step of the transmission process - production of dispersal spores, the distance they travel, and successful colonization of a new host – is strongly mediated by interactions between host and pathogen genotypes. Interestingly, the size of the dispersal unit varies from individual spores to spore clumps, and the size of the dispersal unit has a striking impact on both dispersal distance (Fig. 1) and speed of development of the new infection (Fig. 2).

• Charlotte demonstrated, using both molecular and phenotypic data, that Po. plantaginis is capable of selfing. The production of chasmothecia was also affected by ambient temperature and by the interaction between temperature and pathogen

Fig. 2. The time to sporulation increases as a function of distance from the pathogen source.

8

9

10

11

0 50 100 150Distance from source (cm)

Tim

e to

spo

rula

tion

(day

s)

Fig. 3. Amount of infection in experimental population is affected by both host resistance strategy and co-infection.

genotype. These G, E and G × E may partly explain the high variability in the levels of chasmothecia observed among populations.

• Hanna’s experimental populations of Pl. lanceolata at the Lammi Biological Station demonstrated that disease transmission is strongly affected by the resistance strategy of the host. Also, we find that infection levels are higher, and decline more rapidly, in populations with co-infection compared to populations infected by a single pathogen strain (Fig. 3).

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24 MRG -ANNUAL REPORT 2012

There are more than 250 species of endemic dung beetles in Madagascar in four

large radiations, representing four independent colonizations during the Cenozoic (one clade may date from the late Mesozoic). Since 2002, researchers and students in MRG have constructed molecular phylogenies that include most of the described species, we have sampled beetles across Madagascar to describe their geographical ranges, and we have studied the breeding biology and ecology of selected species and local communities. For a recent review see http://www.mdpi.com/2075-4450/2/2/112.

In the past two years, research has been focused on the radiation consisting of the genera Nanos and Apotolamprus (Canthoninae). This is the youngest of the four major radiations with the estimated age of ca 20 my and 74 described species. Andreia Miraldo has collaborated with Dr Olivier Montreuil (Paris) and Dr. Heidi Viljanen (Helsinki) to revise the taxonomy of Nanos using both genetic and morphological data. We are currently analysing the Nanos-Apotolamprus radiation in detail, examining evolution of body sizes, species’ elevational distributions and their geographical ranges. It is apparent that the radiation started from North Madagascar, from where many

Evolutionary radiations of dung beetles in Madagascar

Undergraduate Kaisa TorppaStudent:

sub-clades have expanded to the west coast and especially to the east coast. The rate of diversification has

generally declined over time, but the emergence of a clade of large-bodied Nanos ca 6 mya is associated with a secondary burst of speciation. These species may have gained competitive advantage because of their large body size. This lineage has expanded its geographical range throughout all of Madagascar and speciated into 24 species mostly in allopatry. We are analysing transcriptomic data for 9 of the 24 species to elucidate the genomic underpinnings of the radiation.

Tanjona Ramiadantsoa is developing a stochastic occupancy model that combines the extinction-speciation dynamics (radiation) with the long-term dynamics of species’ geographical ranges. For the purpose of modelling, Madagascar is divided into five regions within which extinction, ‘sympatric’ speciation and colonization to an unoccupied region are the possible events. If a lineage occurs in more than one region, explicit allopatric speciation is an additional possible event. The model will be fitted to the radiation of the Nanos-Apotolamprus clade using Approximate Bayesian Computation (ABC) methods.

Ilkka HanskiProject leader

Andreia MiraldoPost doc

Anton ChernenkoPost doc

Tanjona Ramiadantsoa PhD -student

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Coevolutionary dynamics and radiations

Collaborators

Olivier Montreuil, the Paris National Museum of Natural History, France

Heidi Viljanen, Finnish Natural History Museum, HelsinkiAri Löytynoja, Institute of Biotechnology, University of

Helsinki.Jukka Sirén, Department of Mathematics and Statistics,

University of Helsinki

Publications

Montreuil O., Viljanen H. and Miraldo A. Phylogeny and taxonomic revision of the genus Nanos Westwood, 1847 (Coleoptera, Scarabaeidae) from Madagascar. In preparation.

Miraldo A. and Hanski I. Tempo and mode of diversification in the radiation of Malagasy dung beetles. In preparation.

Fig. 1. Time-calibrated phylogeny of Nanos-Apotolamprus lineage. Colors indicate the logarithm of body size (length). Body sizes at internal nodes were reconstructed using parsimony in MESQUITE (Alfaro et al. 2009). A significant change in the rate of diversification inferred by MEDUSA is denoted by *. Nodes A and B correspond to nodes where changes in the rate of diversification were detected using the rate of cladogenesis test (Nee et al. 1992). These analyses were done using the apTreeshape (Bortolussi et al. 2006) and GEIGER (Harmon et al. 2008) packages in R.

Miocene Pliocene Pleistocen.

0.02.55.07.510.012.515.017.520.0

Nan.nit.

Ap.amb..

Nan.psem

Nan.con.

Nan.pun.

Nan.dub.

Nan.bic.

Nan.ate.

Ap.ora..

Ap.man..

Ap.ser..

Nan.pey.

Ap.mar..

Nan.mag.

Nan.man.

Ap.zom..

Ap.dar..

Ap.hag..

Nan.han.

Ap.quan.

Ap.han..

Nan.sem.

Nan.and.

Nan.vie4

Nan.vie.

Ap.mil..

Nan.hum.

Ap.cya..

Ap.pse..

Nan.bin.

Nan.mir.

Nan.vad.

Nan.min.

Nan.ran.

Nan.occ.

Nan.pser

Nan.mar.

Ap.per..

Ap.met..

Nan.bim.

Nan.cly.

Ap.sah..

Ap.quam.

Nan.pse.

Ap.pseq.

Nan.rubs

Ap.lat..

Ap.vad..

Nan.mang

Ap.hel..

A

B

* increase in

diversification rate

Ln (Body-size) :

0.92 – 1.17

1.17 – 1.45

1.45 – 1.68

1.68 – 1.93

1.93 – 2.19

2.19 – 2.44 Larg

eb

od

ied

Na

no

s

Fig. 2. Field work in Madagascar. Preparing fish baits for trapping dung beetles (which are mostly attracted by carrion as well), and a plateful of canthonines, mostly Nanos viettei.

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26 MRG -ANNUAL REPORT 201226

Mathematical modelling can be used in ecological research in (at least) two ways. First, mathematical analyses can help to understand

causal relationships. In this context, assumptions are made on the mechanisms behind a phenomenon of interest and mathematical analyses reveal the consequences of these assumptions. Tanjona has taken this approach to analyze what drives spatial community dynamics in heterogeneous landscapes. Second, performing analyses of empirical data often requires advanced mathematical methods. Guillaume, Markku and Jussi are developing various kinds of hierarchical Bayesian approaches with the aim of connecting data to theory. Henjo, Maria, Henna and Ulisses focus on animal movements, with the aim of inferring the underlying mechanisms from the emergent patterns that can be observed in data.

Highlights of the year

In 2012 we welcomed Guillaume and Ulisses as new group members. Having a background in community-level approaches, Guillaume started to implement into an R-package extensions of community-level models

Modelling dispersal and population dynamics

Markku KarhunenPhD -student

Jussi JousimoPhD -student

Maria DelgadoPost doc

Guillaume BlanchetPost doc

Fig. 1. A mathematical framework for analyzing spatio-temporal point processes. Collaboration with mathematicians from Bielefeld University (Yuri Kondratiev and his team) and theoretical ecologists (Stephen Cornell and Ben Bolker) resulted in a mathematical toolbox that applies to a wide range of stochastic and spatial models of interacting individuals.

Tanjona Ramiadantsoa PhD -student

Henjo De KnegtPost doc

Ulisses CamargoPhD -student

Evolution of measures

(model definition, what the individuals do?)

Evolution of spatial moments

(how the model behaves at the population level?)

Evolution of cumulants

(same as above, but a better starting point for analysis)

Henna FabritiusPhD -student

Otso OvaskainenProject leader

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Mathematical ecology

Collaborators

Prof. Juha Merilä, Department of Biological and Environmental Sciences, Helsinki University

Dr. Vesa Selonen and M.Sc. Andrea Santangeli (University of Turku),

Dr. Gonzalo Ferraz, Smithsonian Tropical Research Institute, Manaus, Brazil

Dr. Juan Manuel Morales, Ecotono, INIBIOMA—CONICET, Argentina

Dr. Stephen Cornell, Faculty of Biological Sciences, Leeds University, UK

Prof. Yuri Kondratiev, Dr. Dmitri Finkelshtein and Dr. Oleksander Kutovyi, University of Bielefeld, Germany

Recent Publications

de Boer W.F., van de Koppel S., de Knegt H. J. and Dekker J. J. A. (2013). Hibernation site requirements of bats in man-made hibernacula in a spatial context. Ecological Applications, doi:10.1890/12-0539.1.

Campioni L., Delgado M., Lourenço R., Bastianelli G., Fernandez N., Penteriani V. (2013) Individual and spatio-temporal variations in the home range behaviour of a long-lived, territorial species. Oecologia, doi: 10.1007/s00442-012-2493-7.

Gurarie E. and Ovaskainen O. (2013). Towards a general formalization of encounter rates in ecology. Theoretical Ecology, doi:10.1007/s12080-012-0170-4.

Karhunen M. and Ovaskainen O. (2012). Estimating population-level coancestry coefficients by an admixture F-model. Genetics 192, 609-617.

Saastamoinen M., Brakefield P.M and Ovaskainen, O. (2012). Environmentally induced dispersal-related life history syndrome in the tropical butterfly, Bicyclus anynana. Journal of Evolutionary Biology 25, 2264-2275.

Sundell J., Church C. and Ovaskainen O. (2012). Spatio-temporal patterns of habitat use in voles and shrews modified by density, season, and predators. Journal of Animal Ecology 81, 747-755.

Fig. 2. Field excursion to the Judean desert during a PhD course on animal movement. From the left: Henjo, Otso, Ran Nathan, Luca Giuggioli and Roi Harel.

methods for inferring signatures of natural selection from quantitative trait data, and is currently implementing these methods into an R-package and applying them to different case studies. Henna collected and analysed data on the habitat dynamics and the extinction-colonization

dynamics of the false heath fritillary, to continue with modelling next year.

Otso spent two months of the summer 2012 in Germany, where he participated (as one of the four organizers) the research program “Stochastic Dynamics: Mathematical Theory and Applications” at ZiF (Center for Interdisciplinary Research), Bielefeld. These months were especially important to bring mathematical rigor to the methods we have developed earlier to analyse spatio-temporal point processes (Fig. 1). Tanjona took part to one of the workshops in ZiF with the aim of applying these methods in a model of metacommunity dynamics of wood-decaying fungi.

Otso and Henjo taught in a 5-day PhD Workshop entitled “Movement Ecology: Analyzing Animal Movement Data” at the Hebrew University of Jerusalem. The field excursion to the desert was a hot event (Fig. 2)!

that we have developed earlier. Furthermore, in collaboration with Vesa Selonen and others, Guillaume started modelling the broad-scale occurrence of flying squirrel in Finland. Henjo is also involved in the flying squirrel project but he focuses on dispersal from the point of view of movement analyses. Ulisses started his PhD where his aim is to use a grid of autonomous audio recorders to model the spatio-temporal dynamics of birds in the Amazonian rain forests. This project is conducted in collaboration with Gonzalo Ferraz, whom Otso visited in Manaus last year.

Jussi developed statistical methods for combining individual-level data and population-level data (see also the EBFB project on page 28). Henjo and Maria made progress with analysing the movement behaviour of elephants and eagle owls, respectively. Markku finished the development of statistical

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28 MRG -ANNUAL REPORT 201228

This project (funded by an Academy of Finland research grant for 2011-2015) aims at putting together a broad-scale and long-term database

on European Boreal Forest Biodiversity (EBFB). The EBFB database shall cover the boreal zones of European Russia and Finland and include a description of the environmental changes (e.g. in terms of forest structure and climatic conditions) that have occurred over the past 50 years as well as the population dynamic and phenological responses of various taxonomical groups. The project is based on collaboration between Russian and Finnish participants.

The motivation for compiling the EBFB database is that it makes it possible to address many basic and applied scientific research questions of fundamental relevance for the fields of ecology, conservation biology and climate change research. In particular, we aim to use this database to disentangle the roles of abiotic and biotic factors shaping species abundance, distribution and spatio-temporal population dynamics.

Highlights of the year

In 2012 we organized two workshops, both attended by ca. 20 participants. The first workshop was held

European Boreal Forest Biodiversity (EBFB)

Juri KurhinenProject coordinator

Jussi JousimoPhD -student

Maria DelgadoPost doc

Otso OvaskainenProject leader

at the Mekrijärvi Biological Station and focused on collecting and analyzing winter track data (Fig. 1). The second workshop was a PhD course held in St. Petersburg and focused on mathematical and statistical modelling of ecological data (Fig. 2). In addition, Otso and Juri had a memorable trip to the Komi Republic, visiting the Institute of Biology in Syktyvkar and the Pechoro-Ilychskii Nature Reserve.

In 2012 we compiled a large fraction of the targeted Finnish data (winter track counts on game animals; GPS data on wolf, bear, moose, wild forest reindeer and lynx; small mammal data; many kinds of data on flying squirrel). We acquired pilot versions of Russian data, such as coarse-grained data on winter track counts from European Russia (Fig. 3) and phenology data (e.g. arrival dates of birds, flowering dates of plants, many kinds of weather data for 1960-2012) from the Kivach Nature Reserve. Jussi focused on modelling the spatio-temporal patterns of population abundance whereas Maria took part in the analysis of the phenology data.

IT-designer: Evgeniy MeykeTecnhician: Coong Lo

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Mathematical ecology

Fig. 3. Distributions of wolf (a) and lynx (b) in European Russia in the year 2012. The red dots show observations based on aggregated winter track counts and the colors refer to inferred population density (green=low, orange=high). The maps were produced with the spatial statistics software package INLA-R by Jussi, who conducted a spatio-temporal smoothing utilizing data from 1996 to 2012.

Collaborators

The EBFB project has 26 Russian and Finnish partner organizations, see http://www.helsinki.fi/science/metapop/EBFB/participants.html

In particular, we would like to acknowledge here Prof. Harto Lindén, Dr. Ilpo Kojola and Dr. Jyrki Pusenius (Finnish Game and Fisheries Research Institute), Dr. Ilpo Hanski (Finnish Museum of Natural History), Prof. Andrei Brodsky and Prof. Vladimir Levchenko (St. Petersburg State University), Dr. Marina Yakovleva (Kivach Nature Reserve) prof. Sergei Kochanov and Dr. Eugene Poroshin (Komi Science Centre), Dr. Andrei Kuprijanov (Pechoro-Ilychskii Nature Reserve) and director Anton Bersenev (Hunting and wildlife Ministry of Russia).

Recent Publications

Korpela K., Delgado M., Henttonen H., Korpimäki E., Koskela E., Ovaskainen O., Pietiäinen H., Sundell J., Yoccoz N. G. and Huitu O. (2013). Non-linear effects of climate on boreal rodent dynamics: mild winters do not negate high-amplitude cycles. Global Change Biology, DOI: 10.1111/gcb.12099.

J. Kurhinen, E. Kulebjakina, E. Zadiraka, V.Mamontov, E. Muravskaya, I. Hanski. (2012). Distribution of the Siberian flying squirrel (Pteromys volans L.) in taiga isthmuses between Baltic and White Sea regions. Acta Zoologica Lituanica, 2011, Vol 21(4): 306-310.

Fig. 1. Participants of Mekrijärvi meeting on winter track data.

Fig. 2. Eli Gurarie (who used to be a post doc in our group) taught computer exercises in the PhD course which we organized for Russian PhD students in St. Petersburg.

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Our group develops concepts, theory, algorithms, methods and software for the needs of conservation prioritization – either

spatially or otherwise. We aim at improved ability to put ecologically justified conservation plans and numbers on the political planning table. One part of our work is to understand the factors that explain the distributions of species and communities. We also care about the influence of conservation action on biodiversity features. We then apply optimization, decision theory and uncertainty analysis on top of state-of-the-art ecological modeling, with the aim of identifying efficient and reliable conservation decisions. Major components of our work include species distribution, community and metapopulation modelling, spatial optimization and methods for dealing with spatial connectivity, methods for dealing with habitat loss rates, climate-change considerations, multi-action conservation planning and applications to habitat restoration and offsetting. Understanding the implications of uncertainty to conservation decision making also is relevant for us. We work on applications together with collaborators on all continents.

Atte MoilanenProject leader

Tuuli ToivonenResearcher

Biodiversity conservation informatics

Joona LehtomäkiPhD -student

Undergraduate students: Ninni Mikkonen, Timo Vaahtoranta

Jussi LaitilaResearcher

Federico Montesino PouzolsResearcher

Fig. 1. The cover of the RobOff v1.0 manual – our new software, which is targeted for the allocation of different conservation actions (e.g. habitat management or restoration) and biodiversity offsetting. Cover artwork: Aija Kukkala.

Peter KullbergPhD -student

Enrico Di MininPost doc

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investigations about knowledge gaps in global conservation prioritization, aiming to utilize the GIS data base in a PhD project starting in 2013. Timo Vaahtoranta has been working on an MSc thesis about the classification and taxonomy of conservation strategies, with the intention of soon expanding this work into a PhD thesis.

GEDA members who started last year have been busy with ongoing research. Dr Jussi Laitila has continued his work on the theory and methodological background of spatial ecology and conservation prioritization. His recent work has ranged from the analysis and comparison of conservation planning paradigms to mathematical tools for spatial decision making. Some of his current research interests include the conceptual background of conservation biology, the optimality of conservation decision

making and the analysis of alternative conservation strategies. Dr. Federico Montesino Pouzols has concentrated on developing new methods for spatial conservation planning (within the Zonation

framework) and developing the new RobOff framework and software. He visited ACERA and the University of Melbourne last summer, together with Atte Moilanen, and has been collaborating on various projects in Finland, New Zealand and Australia.

Locally in Finland, the MetZo-project, “Ecologically based decision analysis in the implementation of the South-Central Finland forest biodiversity program”, started in 2010. We develop ecological models for forest and peatland environments together with stakeholders, including much of the Finnish environmental administration. In this project methods developed in our group (Zonation; RobOff) are fitted and applied to conservation decision making in Finland. In this context, Joona Lehtomäki continues working on his PhD thesis on application of spatial prioritization methods to planning of forest conservation and management in Finland, half-time employed by the Finnish Environment Institute. This year, national regional forest analyses were completed and they are being taken onto the field. Also, the national peatland analysis has progressed several rounds and “final” results are expected next spring. The next major objective in this project is the integration of forest and peatland conservation planning.

Also under the MetZO umbrella,Ninni Mikkonen finished her MSc thesis work “Identification of top

Fig. 2. A schematic presentation of spatial prioritization process, which the team attempts to improve either on conceptually or methodologically. Figure adopted from Lehtomäki and Moilanen (submitted).

Datapreparation

Computationalanalysis

Spatial prioritization

Preparation of the ecological model

Preprocesssing of data

InterpretationPriority ranking

Postprocessing

Verification

Setting of objectives

Recommendation for action

”Groundtruthing”

Monitoring

Highlights of the year

This year has been the second of our ERC project GEDA (Global Environmental Decision Analysis). GEDA is funded by a European Research Council (ERC) starting grant, and it provides our base-funding for 2011-2015. GEDA focuses on the development of concepts, analyses, software and large-scale applications of spatial conservation planning.

This year, GEDA and the renewed CoE have brought several new members to our group. Following receipt of a PhD from the University of Kent, Dr Enrico Di Minin started in April with a focus on conservation, economics, and societal impacts of conservation. He has a strong background in modeling, economics and GIS, and a personal interest in conservation in Africa. Dr Tuuli Toivonen (senior lecturer in geo-informatics) started in August. She primarily works on understanding data for spatial conservation and on the combination of land-use zoning and urban planning with biodiversity analyses. Since the spring, Tuuli has been guiding the collation of high-resolution global GIS data for the purpose of spatial modeling and conservation planning. This work has been assisted by Johanna Kuusterä (MSc in geographhy), who is replacing the GEDA research secretary Aija Kukkala during her maternity leave. In the autumn, Peter Kullberg has started preliminary

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by professor Hugh Possingham.Prof. Bill Sutherland and Dr Lynn Dicks of Conservation

Evidence, University of Cambridge.Prof. John Leathwick, Department of Conservation, New

Zealand.Lab of Prof Chris Thomas, University of York.

Recent key publicationsPouzols F.M., Burgman M.A., and Moilanen A. (2012).

Methods for allocation of habitat management, maintenance, restoration and offsetting, when conservation actions have uncertain consequences. Biological Conservation, 153: 41-50.

Laitila J. and A. Moilanen. (2012). Use of many low-level conservation targets reduces high-level conservation performance. Ecological Modelling 247: 40-47.

Kujala H., Burgman M.A., and Moilanen A. (2013). Treatment of uncertainty in conservation under climate change. Conservation Letters, early doi: 10.1111/j.1755-263X.2012.00299.x.

Kukkala A. and A. Moilanen. (2013). The core concepts of spatial prioritization in systematic conservation planning. Biological Reviews, doi: 10.1111/brv.12008.

CollaboratorsNational

The South-Central Finland Forest Biodiversity Programme (METSO) and its ecological decision analysis umbrella project, MetZo, chaired by Dr Mikko Kuusinen at the Ministry of Environment. The MetZo project involves the Finnish Ministry of Environment, the Ministry of Agriculture and Forestry, the Finnish Environment Institute, the Forest and Park Service (Metsähallitus), the Finnish Forest Research Institute, universities, forestry centers, environmental centers (ELY).

The NatNet LIFE+ project for improving the conservation area network of South-Western Lapland. (involves e.g. the Forest Research Institute, Environment center & Forestry center Rovaniemi / Lapland)

Prof. Janne Kotiaho, Prof. Mikko Mönkkönen, Dr Adriano Mazziotta, and Mr Santtu Kareksela, University of Jyväskylä.

InternationalACERA, Australian Centre of Excellence in Risk Analysis,

directed by professor Mark BurgmanCEED, Australian Centre of Excellence in Environmental

Decisions, University of Queensland, Australia, directed

priority areas and management landscapes from a national Natura 2000 network”. In this work, a prioritization was developed across the Finnish Natura 2000 network, with applications for the targeting of habitat protection, maintenance and restoration. Currently, Ninni works at the Natural Heritage Services of Metsähallitus (Finnish forest and park service) to help decide expansions of the Finnish forest protection area network.

The group leader Atte Moilanen has mainly worked with initiation of new projects together with the expanded group. Some topics presently of interest for him include the concept of conservation value, methods for habitat management and restoration, development of new operational approaches for multi-action conservation planning, development of methods and analyses relevant for the Nagoya CBD resolution, and analysis of alternative conservation strategies. This year, Atte Moilanen together with Jussi Laitila and Federico Montesino Pouzols taught two courses “Concepts and principles of spatial conservation planning”, and “Methods and software of spatial conservation prioritization”, both 3 ECTS.

Our oldest software product, Zonation (since 2004), has improved a lot this year. Dr Federico Montesino Pouzols has implemented great improvements into both the core and user interface of the Software. The V3.1 manual was printed and distributed spring 2012, and presently we are working towards version 4.0. During 2012, Victoria Veach has put in major effort reviewing and revising the documentation of Zonation 4 as well as other documents, including www pages, teaching material, etc.

Next year we expect (i) to continue the development of software Zonation and Roboff, (ii) the completion of development of the global high-resolution GIS data base for global analyses of conservation priority , (iii) initiation of a major new project for the analysis of conservation strategies, which was 12/2012 kindly funded by the Kone Foundation (iv), progress for both forests and peatlands at the national planning stage, (v) expansion of the group by two PhD students (Peter Kullberg, Timo Vaahtoranta and Victoria Veach) to formally start during the spring.

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Conservation effectiveness

Mar CabezaProject leader

Undergraduate student: Helena Uotila

Johanna Eklund PhD -student

Protected areas cover twelve percent of the Earth’s terrestrial surface, yet they are insufficient. They are inefficient in covering biodiversity, and they

are ineffective in protecting it. Population declines as well as illegal deforestation in protected areas worldwide indicate that reserves are not functioning. Part of the problem stems from a history of ad-hoc decisions, which has motivated advances in the science of reserve-network design. Nevertheless, a substantial gulf remains between what is moving research versus what practitioners claim to require. Our work aims at assessing the effectiveness of conservation actions, including protected areas and management outside them. We also strive to understand potential causes for reduced effectiveness. Our projects expand from large scale, global analyses to very local empirical studies. Examples from larger to smaller scale assessments include: Evaluating the role of protected areas in reducing tropical deforestation; Appraising EU’s solidarity funds from the biodiversity conservation perspective; Assessing the effectiveness of Finnish conservation policies in relation to the Red List Index; Examining the effectiveness of local management practices on the population viability of a butterfly in Finland.

Fig. 1. Investigating deforestation causes in Madagascar: Resource mapping activity at a Malagasy village.

Silvija BudaviciutePhD -student

Henna FabritiusPhD -student

Anni ArponenPost doc

Ricardo RochaPhD -student

Laure ZupanPhD -student

Highlights of the year

Johanna visited Lauren Coad’s team (Oxford) to discuss the deforestation project and future collaboration in investigating the links between governance and effectiveness. Ricardo and Johanna have been analyzing deforestation in Madagascar and

© Helena Uotila

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Helena spent three months in the field conducting interviews, to try to understand the social drivers of some of the patterns encountered (Fig. 1.). The visit of Mr. Mamy Rakotoarijaoana, the Ranomafana National Park director (Madagascar) initiated an interesting collaboration from which we expect new research angles. Silvija has concentrated on sorting and identifying the massive ant samples she got from Madagascar in late 2011, and has also measured morphological characters of over 2000 individuals. These will be used to address the importance of functional traits and trait groups in performing particular ecosystem functions in disturbed habitas. Ricardo has now completed one and a half years of fieldwork in Central Amazon, Brazil. His PhD aims at assessing the long-term impacts of deforestation upon tropical biodiversity, using bats as a study model (Fig. 2.). Laure has started to evaluate a facet of biodiversity that is often ignored in effectiveness evaluations: the representation of evolutionary history in conservation areas. She found that within Europe, protected areas fail to cover important areas of high phylogenetic diversity. Finnish conservation policies are also receiving more of our attention. What started as a paper discussion in our regular journal club, has turned into a rather large collaborative project. Anni is leading this collaboration, which looks into the impact of conservation investments on trends in extinction risk. Thus far we have put substantial effort into collecting comprehensive datasets, which we are now beginning to analyse. Anni is also leading a project on the effectiveness of Finnish policies in

Fig. 2. Pteronutus personatus, one of the new species added to the Biological Dynamics of Forests Fragments Project (BDFFP), in Central Amazon, Brazil , during Ricardo’s survey in the area this year.

Collaborators

Lauren Coad, ECI, University of Oxford, UKNeil Burgess, WWF USAMamy Rakotoarijaona, Madagascar National ParksWilfried Thuiller, CNRS, FranceTarmo Virtanen, Dept Env Sciences, University of HelsinkiAino Juslén, Andrea Santangeli, Finnish Museum of

Natural HistoryMikko Kuussaari, Saija Sirkiä Finnish Environment

Institute, Natural Environment Centre Riikka Paloniemi, Jukka Similä, Finnish Environment

Institute, Environment Policy Centre

Publications

Arponen A., et al. Improving conservation planning of semi-natural grasslands: integrating connectivity into agrienvironment schemes. Biological conservation, under revision.

Zupan L. Cabeza M., et al. A conservation dilemma: spatial mismatch of phylogenetic diversity across three vertebrates groups and protected areas in Europe. Proceedings of the Royal Society B., under revision.

conserving farmland biodiversity in collaboration with the Finnish Environment institute. Henna has collected and analyzed institutional and ecological data to demonstrate the deficiency of short-term management plans for her study species, the endangered butterfly M. diamina.

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Mar CabezaProject leader

Climate change

Undergraduate student: Antti Takolander

Our team covers a broad range of topics in relation to climate change and biodiversity. Our research interests range from methods to

assess the observed impacts on species and conservation areas, to developing and improving tools to project future impacts. Furthermore we develop approaches to include future projections in conservation planning. We are interested in addressing the plethora of uncertainties at various levels, in particular those related to predicting impacts and decision-making processes. We acknowledge important indirect impacts of climate change on biodiversity that result from mitigation and adaptation responses in other societal sectors. We thus work closely with researchers covering other fields (energy, agriculture, water, health) in order to identify potential positive and negative interactions.

Highlights of the year

Heini and Maria, successfully defended their PhD thesis this year. They have also been successful in securing postdoc positions (Heini at the University of Melbourne; Maria at the University of Jyväskylä) and continue collaborating with us. Heini’s recent findings relate largely to uncertainty in climate change research and have resulted in three papers. She investigated how spatiotemporal changes in survey effort affect measurements of observed range shifts

Fig. 1. Whinchat, Saxicola rubetra, one of the species projected to be most exposed and most sensitive to global.change in the Iberian Peninsula (Triviño 2012).

Maria TriviñoPhd -studentRaquel Garcia

PhD -student

when using atlas data. She also reviewed how different types of uncertainties are addressed in climate change research, and demonstrated that despite uncertainties, scientifically robust solutions both in modelling of future impacts and in spatial conservation prioritization can be achieved. Maria showed that climate remains the most important factor for modelling the distribution of species in the Iberian Peninsula. She also found that the species most exposed to future climate changes are not those with most sensitive to it, according to their

Heini KujalaPhd -student

Laura MellerPhD -student

Astrid van TeeffelenPost doc

© Ricardo Rocha

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traits and threat status, with few exceptions (Figure 1). Raquel’s work has focused on climate change impacts on African vertebrates She has investigated the methodological uncertainties in results from species distribution models (SDM). She found most of the variation in projections to result from the choice of modelling algorithm, and compared different ways to generate consensus around the projections (Figure 2). She is currently looking at whether species at risk identified by SDMS match with intrinsic traits that potentially influence the species’ response.

Much of the work produced sits within the EU-FP7 project RESPONSES (2010-2013), for which Mar leads the Biodiversity work package. RESPONSES investigates EU policy action on climate change covering a range of sectors and communicating findings to the EU. In August, Mar gave a well-received keynote speech at the Nordic Climate Adaptation conference, summarizing the outcomes of RESPONSES project. We also produced a policy brief that stressed the gaps between science and policy and also highlighted the expected reduction in European conservation effectiveness due to climate change. We organized a workshop with biodiversity experts in Ehrenberg, Germany to assess cross-sectoral opportunities and threats from mitigation and adaptation. The outcomes are being combined with a literature review and a model analysis. Laura is preparing a manuscript exploring the relative impacts of climate change and bioenergy, thereby addressing the

Fig. 2. Maps illustrating the major sources of uncertainties in projections of species turnover for the year 2050. Variation is larger among different Bioclimatic Envelope Models (BEM) than emissions scenarios (SRES), although some locations in Africa also show disagreement between climate models (GCMs) (see Garcia et al. 2012)

Collaborators

Miguel B. Araújo (National Museum of Natural Sciences, Spain)

Neil Burgess, WWF US Mark. A. Burgman, University of MelbourneJon E. Brommer, University of TurkuSebastiaan Deetman, Netherlands Environment Assessment

AgencyWendy Foden, IUCN, Cambridge, UKThomas Hickler, Goethe University, GermanyAndries Hof, Netherlands Environment Assessment AgencyAleksi Lehikoinen, Finnish Museum of Natural HistoryCarsten Rahbek, University of Copenhagen , DenmarkWilfried Thuiller, University of Grenoble, France

Key publications

Garcia, R. A., N. D. Burgess, M. Cabeza, C. Rahbek, and M. B. Araújo. (2012). Exploring consensus in 21st century projections of climatically suitable areas for African vertebrates. Global Change Biology 18:1253–1269

Kujala, H. (2012). Climate change, species range shifts and uncertainty – a new era of conservation planning. PhD thesis, Faculty of Biological and Environmental Sciences, University of Helsinki. Unigrafia Oy, Helsinki.

Kujala, H., Burgman, M.A. and Moilanen, A. Treatment of uncertainty in conservation under climate change. (2013). Conservation Letters, doi: 10.1111/j.1755-1263X.2012.00299.x

Kujala, H., Moilanen, A., Araújo, M.B. and Cabeza, M. (2012). Conservation planning with uncertain climate change projections. PLOS ONE (in press)

Kujala, H., et al. (2013) Range margin shifts of birds revisited — the role of spatiotemporally varying survey effort. Global Change Biology, 19:420-430.

Meller, L., Barbet-Massin, M., Deetman, S., Hof, A., Pironon, S., Thuiller, W., & Cabeza, M. (2012). Joint impacts of climate change and bioenergy production on the state and conservation of European birds. RESPONSES project deliverable 5.3a (p. 30 pp.).

Triviño, M., 2012. Global change impacts and conservation priorities in the Iberian Peninsula. PhD dissertation in Ecology. University Rey Juan Carlos and National Museum of Natural Sciences, Madrid.

Van Teeffelen, A., Lung, T., Meller, L., Vermaat, J., Lavalle, C., & Cabeza, M. (2012). EU Biodiversity policy in times of climate change. RESPONSES policy update No. 1.

Van Teeffelen, A.J.A., Vos, C.C. & Opdam, P. (2012). Species in a dynamic world: Consequences of habitat network dynamics and conservation planning. Biological Conservation, 153, 239-253.

mitigation versus adaptation debate. Antti Takolander started his MSc with us, comparing projections from mechanistic models to those of correlative species distribution models, for a selected number of tree species.

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Supporting personnel

Suvi IkonenCoordinator

My main task is to take care of the Glanville fritillary (Melitaea cinxia)

laboratory populations and assist researchers in their field and laboratory experiments. I’m working year around at Lammi Biological Station.

I am taking care of data recording during the Glanville fritillary survey and data

extraction from the database for different projects. I am also programming user interface and data handling components for software produced in the group.

Evgeniy MeykeIT designer

I facilitate the laboratory work of the molecular ecology team by

providing the necessary materials, chemicals and enzymes. Extracting, amplifying and sequencing DNA is an essential part of my tasks as well as processing samples for screening of DNA micro satellites. I also process samples for SNP (single nucleotide polymorphism).

Toshka NymanLaboratory technician

My tasks in the Glanville fritillary project include practical lab work and also

planning and testing laboratory methods. I prepare samples for high-throughput genotyping. My work includes mainly sequence comparisons, primer design and PCR-testing before large scale genetic analyses.

Annukka RuokolainenLaboratory technician

I do the laboratory work in the project of wood-decaying fungi. Mainly that includes extracting

DNA from wood and spore samples and processing them for 454-sequencing. I also work for the faculty in MES laboratory, assisting other groups in their lab work.

Heini Ali-KoveroLaboratory technician

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Supporting personnel

My main duties consist of group’s financial and personnel administration as well as international project manage-

ment. I am also taking care of various other research supporting tasks.

Viia ForsblomResearch secretary I am organizing and supervising large

scale annual surveys of Glanville fritillary butterfly in the Åland islands every spring and fall. In the meantime I’m helping people with all kinds of IT and other technical tasks in the group.

Sami OjanenProject coordinator

Marja-Leena PeltonenLaboratory technician

I am a lab technician in Mikko Frilander’s group. My main responsibilities are RNA and DNA isolations from butterfly,

beetle and other biological materials, PCR, RT-PCR, cloning etc.I also take care of reagent supplies in Mikko’s lab.

Suvi SaarnioLaboratory technician

I do laboratory work in the Glanville fritillary project. For now my main tasks in the lab are performing GoldenGate

Genotyping Asseys and preparing samples to other high throughput genotyping analyses. My duties also include testing new laboratory methods and improving protocols.

My main tasks in the Biodiversity Con-servation Informatics Group include

general administration, project management, reporting and organization of events. I’m also helping with miscellaneous research-related tasks, preparation of teaching materials and documentation of software. On maternity leave.Aija Kukkala

Research secretary

I I work in Anna-Liisa Laine�s group of species interactions in metapopulations.

My tasks are divided between office, greenhouse and laboratory. I also coordinate most of the practical issues of mildew surveys in Åland. In other words I am secretary, gardener, powdery mildew babysitter and project coordinator in a same person.

Krista RavealaResearch secretary

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Pia VälitaloTechnical assistant

I do laboratory work in the Glanville fritillary project. I am in charge of DNA and RNA sample management and Progeny

database management. In addition I’m doing most essential lab work, which includes DNA extraction, gel analysis, PCR, DNA quantification and qualification and so on.

Victoria VeachTechnical assistant

I have been editing the Zonation and Roboff manuals for Atte. I have also been helping

with data pre-processing for Mar’s group.

Johanna KuusteräResearch secretary

I am taking care of personnel and financial administration tasks and working with GIS

database building and other tasks.

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Synopsis of the year 2012

Publications

ThesesArticles and book chapters

Honours, awards and memberships

Visitors

Teaching

Budget

Annual meeting in St Petersburg

©Sami Ojanen

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Heini Kujala , PhD (June 1st): Climate change, species range shifts and uncertainty -A new era of conservation planning

PhD theses

All species are adapted to certain climatic conditions, outside of which they cannot survive. Changes

in the climatic environment therefore force species to either adapt to the new conditions or move to areas where suitable conditions are still present in order to avoid extinction. Several studies have shown that species from various taxa are currently moving their ranges polewards and to higher elevations to keep up with shifting climate regimes. However, species differ widely in their dispersal abilities. In addition, natural landscapes are becoming increasingly human-dominated, further hindering dispersal by decreasing permeability. anthropogenic climate change is therefore expected to become one of the major drivers of species extinctions by the end of the 21st century.

Species range shifts are problematic in conservation planning, because dynamic biodiversity patterns hamper our ability to identify priority areas for protection. because protected area networks are geographically fixed, climate change may also drive species out of reserves, foiling past conservation efforts. in this thesis the different risks and opportunities of conducting conservation planning under climate change are investigated. This research focuses on the uncertainties that arise from working with unknown future events, technical challenges of observing and predicting species range shifts, and using (or ignoring) information about future impacts in conservation planning.

The major findings of this thesis are that climate change is already rapidly reshaping species distributions in Finland and that ignoring future

dynamics can lead to misguided and potentially inefficient conservation decisions. The results presented here show that modelling future impacts using so-called niche modelling techniques, despite their inherent uncertainties, can provide useful information about how species distributions and conservation statuses will be affected by climate change. For example previously created models for Finnish breeding birds predicted well recently observed changes in species distribution sizes. More importantly, the observed changes seem to match best with predictions that follow the most extremeclimate change scenario. A key factor for successfully measuring and predicting climate change impacts are good monitoring data, the role of which should be more widely acknowledged by decision-makers. uncertainty in climate change research is pervasive and cannot ever be entirely eliminated. This work offers tools to assist in both spatial prioritization and decision making when scarce conservation resources need to be allocated under uncertain future conditions. The findings of this thesis strongly encourage using proactive approaches that account for future impacts. The results also suggests that while striving to reduce epistemic uncertainty is important in climate change and conservation research, other sources of uncertainty such as socio-political factors or volitional human behaviour might constitute far larger determinants of successful conservation actions, and therefore merit stronger focus in research.

Supervisor: Mar Cabeza

Letizia Campioni , PhD (University of Seville) (February): Breeders and nonterritorial individuals of a Long-lived species, the Eagle owl Bubo bubo: Differences in space use and movement patterns

Many animal species live in societies in which nearby conspecifics are vital elements of their

social environment, with the nature and quality of their behavioral interactions determining the type of social organization. As a group, birds show a wide range of social organizations where, in some cases, social status gives priority of access to resources, ultimately affecting

individual fitness. For example, in territorial species where at least two social groups –breeders and non-territorial floaters – are recognized, territorial ownership can lead to holders behaving differently compared to the floating counterpart of the population. For this reason, social structure is often considered a key determinant of population biology, influencing fitness, gene flow, and

Publications

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Synopsis of the year

MSc theses

Riikka Alanen, MSc: Effects of habitat fragmentation on fitness and genetic variability of wind-pollinated plant species Plantago lanceolata

In my MSc thesis I studied the effects of habitat fragmentation on fitness and genetic diversity in

a wind-pollinated plant species Plantago lanceolata. Human induced habitat fragmentation is one of the most severe problems for survival of many species. Habitat fragmentation reduces population sizes and isolates populations from each other. In smaller and isolated populations the risk of inbreeding depression increases and that affects fitness of the populations. The effects of habitat fragmentation on plant fitness and genetic variation may also depend on plant characteristics such as life span, mating system and rarity.

The experiment was carried out in 2010-2012 and it was part of a more extensive study which aims to find out how well different proxies account for plant’s life-time reproduction success. 40 populations representing the natural variation of the system in Åland Islands were chosen for the study. The study species Plantago lanceolata is a diploid perennial. It is self-sterile and wind-pollinated plant that also proliferates clonally. It has a tendency of forming a soil seed bank where the seeds may survive viable for a number of years. The population in Åland Islands is naturally fragmented and located at the Northern edge of this species range .

spatial pattern and scale. Nonetheless, nonsocial factors (e.g., environmental condition, food supply) can also affect behavioral interactions, individual relationships, and, ultimately, social organization. In the present thesis, we studied the behavioral differences between individuals of different social status; we focused in particular on the analysis of habitat selection, space use behavior and movement patterns of breeders and nonterritorial eagle owls (Bubo bubo).

The focal radio-tracking of breeders and nonterriotrial floaters during 8 years demonstrated that owls perform different behavioral strategies in relation to different life cycle stages, social status and the behavioral trait under study. These observations emphasize the existence of more structured inter-individual relationships than expected. Moreover, previous investigations of social interactions (vocal and visual communication) support the importance of territoriality and social dominance on owls’ behavioral decisions. Our results indicate a scenario in which both social and nonsocial factors seem to affect the behavioral mechanisms that regulate habitat selection, space use and movement behavior in different ecological contexts. In contrast to our initial predictions, trophic resource abundance in our study area does not correlate directly with owls’ space use behavior. However, the large abundance of the staple prey across this area, due primarily to management and release of rabbits (the study area serving as game reserve), might actually favor a high density of conspecifics over a reduced area (40 breeding pairs/100 km2) by relaxing environmental constrictions like resource competition (e.g. food). In line with this prediction, we show that territory holders (Chapter 5) occupy reduced home ranges of high quality for reproduction. Surprisingly,

the home range size (mean HR size ~ 220 ha) appears to be a direct consequence of landscape structure rather than prey abundance available across the study area. Across the mosaic of territories settled by owls, females - the sex that experiences less social constriction - are those which exhibit wider home ranges that overlap to a greater extent with those of their neighbors. Nevertheless, within the boundaries of their home ranges, adults’ behavioral decisions were significantly affected by nonsocial factors such as the biological needs and individual identity. Similarly, external cues like the lunar cycle (Chapter 3), act to regulate the time and effort that owls allocated to social (communication) or physiological (feeding) activities.

A key finding demonstrated here is that nonterritorial floaters show a tremendous capacity to adapt their behaviour to their immediate needs and social and physical surroundings. As with other territorial species, floating owls show cryptic behaviour, living in a parallel “underworld” where individuals make decisions while considering social constraints (Chapter 1-2), acquired experience (Chapter 4) and landscape features. At the end of their natal dispersal, the most likely fate for our floaters was to settle close to the natal population while awaiting circumstances that would offer greater reproductive opportunities. In conclusion, the study of the relationships between animals and their environment is a field where ecology and behavior are tightly intertwined. In my opinion, and as stressed in the present study, social organization is a key determinant of population biology with important implications on spatial processes.

Co-Supervisor: Maria Delgado

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Peter Kullberg, MSc: Habitat Fragmentation and the Performance of Plantago lanceolata and Veronica spicata

Phenotypes of both species showed some sensitivity to the size and isolation levels of local populations, but overall the detected effects were relatively small. Individuals from small local population of P. lanceolata were on average smaller than individual from large populations (F(1, 43.2) = 5.04, p = 0.031). Leaf size of V. spicata was smaller in highly connected populations (F(1, 33.6) = 9.27 p = 0.005). The effects of populations size to leaf size of V. spicata was different within stress groups (Interaction: F(1,117) = 4.04, p = 0.029). In stressed group leaf size was largest in small populations, but in non-stressed leaf size was largest in large populations.

None of the detected effects could be unambiguously accounted to be caused by inbreeding depression. The effects were more likely caused by environmental factors that are correlated with population isolation and size. Larger size of V. spicata in isolated populations could also be caused by adaptation to pollen limitation in isolated populations. This study has shown that fragmentation does not necessarily lead to inbreeding depression in small and isolated populations and species can be adapted to living in fragmented conditions. Interpopulation gene flow, purgin, and certain species characters can be responsible for the low levels of inbreeding depression.

Supervisors: Anna-Liisa Laine Saskya van Nouhuys

Habitat fragmentation is one of the major threats to species diversity around the world. It increases the

distance between remnant habitat patches and decreases their size. As a result, the population sizes of the species inhabiting the fragments is reduced and their isolation is increased. Isolation and small population size can lead to inbreeding and loss of genetic variation. Inbreeding depression is expected to cause reductions in the fitness traits of the species inhabiting the remnant fragments. Habitat fragmentation alters also the environmental conditions and species interactions of the populations living in remnant patches. This can lead to changes in the selection pressure of the fitness traits and further to adaptive evolution.

In this thesis, the effects of habitat fragmentation on two plant species, wind pollinated P. lanceolata and insect pollinated V. spicata, are studied with a common garden experiment. Both species are perennial herbs living in fragmented environment in South-West Finland. From both species, four individuals from 40 different populations with different size and isolation levels were used. Two treatments stress and reduced pollination were applied. In the end of growing season fitness related traits from the plants were measured. Principal components were extracted based on the measurements. Components of P. lanceolata were: plant size, number of flowers and leaves, and see set. Components of V. spicata were: leaf size and number of leaves. Effects of isolation and population size to the principal components were analysed with linear mixed models.

I studied if the degree of isolation and size of the population have any effect on seed weight, germination and plant size. The only significant correlation was between population connectivity and plant size, plants in isolated populations were smaller than in populations that were better connected to other populations.

AMOVA analysis of the genetic data revealed that the amount of inbreeding in these populations is fairly high. Still the populations did seem to have some gene flow among them, because the populations didn’t seem to be strongly differentiated from each other. The inbreeding didn’t seem to affect viability of the seeds (germination percentage being roughly at the same level as in continuous populations. Plantago lanceolata

populations in Åland Islands don’t seem to be seriously affected by the fragmented population structure and the reason for that may be found from the plant characteristics such as clonal reproduction and the formation of extensive soil seed bank.

The state of Plantago lanceolata in Åland seem to be fairly good at the moment, but it is advisable to keep track on the state of the population in future because the negative impacts of inbreeding may arise if the population suffers from stressful conditions.

Supervisors: Anna-Liisa LaineSaskya van Nouhuys

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Synopsis of the year

The purpose of this master’s thesis is to study nature values within the Finnish national Natura 2000 network on state owned land. The six goals of this work were achieved: 1) Areas with most nature values were identified by prioritizing habitats of Natura 2000 directive (92/43/EU) within. Areas with high nature value were usually in very natural state and had good connectivity to other similar places, or they were spots of some very rare nature types. 2) It was found out that data used was suitable for identifying conservation values, 3) find out the suitability of Zonation software in conservation area management and maintenance planning and 4) find out how results will change if conservation status is taken into account. As an addition to these 5) the most considerable areas with high conservation value were identified and 6) “Zonation software in a nutshell” was produced in Finnish to assist Finnish state officials to use the software for conservation purposes. These results will help Metsähallitus (The Finnish Forest and Park Service) - Natural Heritage Service - to target resourcing of habitat management and restoration in and around the areas with most considerable nature values. It is essential to sustain these areas and their values so that their nationwide importance can be maintained into the future.

Data used in this study covered areas that were classified as Natura 2000 habitats according to European Union Council Directive 92/43/EEC. Analyses were done by using Zonation software, a tool for spatial

Ninni Mikkonen, MSc: Most valuable areas of Natura 2000 habitat types in Finnish conservation areas

conservation prioritization. Data consisted of 68 Natura 2000 habitat types and their state of naturalness and representativeness. Zonation took into account the rarity, quality, importance, threat status, biodiversity value, congruity and connectivity of these habitat types. As a result software produces a map of conservation priorities and associated quantitative information, which facilitate identification of areas with most considerable nature values. These were identified both ocularly and with Zonation software.

Analyses were done at two levels: all habitat types together and in subgroups following division to major habitat types, such as coastal environments, inland waters, meadows, alpine habitats, peat lands, rocky areas and forests. Results showed that connectivity increased aggregation of areas with high nature values and weighting spread them. Hierarchical analysis was used to find out how nature values changed when the conservation status of the areas were taken into account. The results of hierarchical analysis show that conservation status changed the results a lot. Difference between main analysis and hierarchical analysis was much greater than when taken into account connectivity of feature weights. Hierarchical comparison revealed that many areas with considerable high nature values areas are not presently strictly protected.

Supervisor: Atte Moilanen

Articles and book chapters

Alagador D. A., Triviño M., Cerdeira O., Bras R., Cabeza M., and Araújo M. B. (2012). Linking like with like: optimising connectivity between environmentally-similar habitats. Landsc. Ecol. 27:291-301.

Arponen A. (2012). Prioritizing species for conservation planning. Biodivers. Conserv. 21:875-893.

Arponen A., Heikkinen R. K., Paloniemi R., Pöyry J., Similä J., and Kuussaari M. (in press). Improving conservation planning for semi-natural grasslands: integrating connectivity into agri-environment schemes. Biol. Conserv.

Arponen A., Lehtomäki J., Leppänen J., Tomppo E., and Moilanen A. (2012). Effects of connectivity and spatial resolution of analyses on conservation prioritization across large extents. Conserv. Biol. 26:294-304.

Bekessy S. A., White M., Gordon A., Moilanen A., McCarthy M. A., and Wintle B. A. (2012). Transparent

planning for biodiversity and development in the urban fringed. Landsc. Urban Plann. 108:140-149.

Bonte D. and Saastamoinen M. (2012). Dispersal syndromes in arthropods: butterflies and spiders as a model system. in M. Baguette, T. Benton, J. Bullock, and J. Clobert, editors. Dispersal and Spatial Evolutionary Ecology. Oxford University Press, Oxford.

Bonte D., Van Dyck H., Bullock J. M., Coulon A., Delgado M. d. M., Gibbs M., Lehouck V., Matthysen E., Mustin K., Saastamoinen M. A. K., Schtickzelle N., Stevens V. M., Vandewoestijne S., Baguette M., Barton K., Benton T. G., Chaput-Bardy A., Clobert J., Dytham C., Hovestadt T., Meier C. M., Palmer S. C. F., Turlure C., and Travis J. M. J. (2012). Costs of dispersal. Biological Reviews. 87:290-312.

Calandra R. R. T., Deisenroth M. P. & Pouzols F. M. 2012. Learning Deep Belief Networks from Non-stationary Streams. Pages 379-386 in Artificial Neural Networks and Machine Learning – ICANN 2012. Springer Berlin Heidelberg.

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Campioni L., Lourenco R., Delgado M. d. M., and Penteriani V. (2012). Breeders and floaters use different habitat cover: should habitat use be a social status-dependent strategy? Journal of Ornithology. 153:1215-1223.

Campioni L. D. M., Lourenço R., Bastianelli G., Fernandez N., Penteriani V. (2013). Individual and spatio-temporal variations in the home range behaviour of a long-lived, territorial species. Oecologia. doi: 10.1007/s00442-00012-02493-00447.

de Boer W. F., van de Koppel S., de Knegt H. J., and Dekker J. J. A. (2013). Hibernation site requirements of bats in man-made hibernacula in a spatial context. Ecol. Appl.: doi:10.1890/1812-0539.1891.

De Jong M., Collins S., Beldade P., Brakefield P., and Zwaan B. (2013). Footprints of selection in wild populations of Bicyclus anynana along a latitudinal cline. Mol. Ecol. 22:341-353.

Dees M. W., Somervuo P. J., Lysøe E., Aittamaa M., and Valkonen J. (2012). Species identification and microarray-based comparative genome analysis of Streptomyces species isolated from potato scab lesions in Norway. Mol. Plant Pathol. 13:174-186.

Di Minin E., Fraser I., Slotow R., and MacMillan D. C. (2013). Understanding heterogeneous preference of tourists for big game species: implications for conservation and management. Anim. Cons.: DOI: 10.1111/j.1469-1795.2012.00595.x.

Di Minin E., MacMillan D. C., Goodman P. S., Escott B., Slotow R., and Moilanen A. (2013). Conservation businesses and conservation planning in a biodiversity hotspot. Conserv. Biol.

Di Minin E. and Moilanen A. (2012). Empirical evidence for reduced protection levels across biodiversity features from target-based conservation planning. Biol. Conserv. 153:187-191.

Duplouy A., Iturbe-Ormaetxe I., Beatson S. A., Szubert J. M., Brownlie J. C., McMeniman C. J., McGraw E. A., Hurst G. D., Charlat S., O’Neill S. L., and Woolfit M. (2013). Draft genome sequence of the male-killing Wolbachia strain wBol1 reveals recent horizontal gene transfers from diverse sources. BMC Genomics. 14:20.

Garcia R. A., Burgess N. D., Cabeza M., Rahbek C., and Araujo M. B. (2012). Exploring consensus in 21st century projections of climatically suitable areas for African vertebrates. Global Change Biology. 18:1253-1269.

Gurarie E. and Ovaskainen O. (2013). Towards a general formalization of encounter rates in ecology. Theoretical Ecology. doi:10.1007/s12080-12012-10170-12084.

Halme P., Vartija N., Salmela J., Penttinen J., and Norros V. (2012). High within- and between-log variation in the nematoceran community and its physical environment in decaying aspen trunks. Insect Conservation and Diversity.

Hanski I. (2012). Dispersal and eco-evolutionary dynamics in the Glanville fritillary butterfly. Pages 290-303 in J. Clobert, Baguette, M., Benton, T.G. and Bullock, J.M., editor. Dispersal Ecology and Evolution. Oxford University Press, Oxford.

Hanski I. (2012). Eco-evolutionary dynamics in a changing world. Year in Ecology and Conservation Biology. 1249:1-17.

Hanski I. (2012). Eco-evolutionary dynamics in a changing world. Pages 1-17 in R. S. Ostfeld and W. H. Schlesinger, editors. The Year in Ecology and Conservation Biology. Annals of the New York Academy of Sciences.

Hanski I. (2012). Metapopulations and spatial population processes. Oxford Bibliographies in Ecology. http://www.oxfordbibliographies.com/.

Hanski I. (2012). Spatial structure and dynamics in the Glanville fritillary (Melitaea cinxia) metapopulation. Pages xxviiii-xxxi in J. Clobert, Baguette, M., Benton, T.G. and Bullock, J.M. , editor. Dispersal Ecology and Evolution. Oxford University Press, Oxford.

Hanski I. (2012). A tribute to the ERC-long live basic research. EMBO Rep. 13:474-474.

Hanski I. and Kuuluvainen T. (2012). Metsien monimuotoisuuden väheneminen jatkuu. Helsingin Sanomat.

Hanski I., von Hertzen L., Fyhrquist N., Koskinen K., Torppa K., Laatikainen T., Karisola P., Auvinen P., Paulin L., Makela M. J., Vartiainen E., Kosunen T. U., Alenius H., and Haahtela T. (2012). Environmental biodiversity, human microbiota, and allergy are interrelated. Proc. Natl. Acad. Sci. U. S. A. 109:8334-8339.

Hornett E. A. and Wheat C. W. (2012). Quantitative RNA-Seq analysis in non-model species: assessing transcriptome assemblies as a scaffold and the utility of evolutionary divergent genomic reference species. BMC Genomics. 13:361.

Karhunen M. and Ovaskainen O. (2012). Estimating Population-Level Coancestry Coefficients by an Admixture F Model. Genetics. 192:609-617.

Karinen S. H., Saarinen S., Lehtonen R. J., Rastas P., Vahteristo P. M., Aaltonen L. A., and Hautaniemi S. (2012). Rule-based induction method for haplotype comparison and identification of candidate disease loci. Genome Med. 4:21.

Kool J. T., Moilanen A., and Treml E. A. (in press). Population connectivity: recent advances and new perspectives. Landsc. Ecol.: doi: 10.1007/s10980-10012-19819-z.

Korpela K. D. M., Henttonen H., Korpimäki E., Koskela E., Ovaskainen O., Pietiäinen H., Sundell J., Yoccoz N. G. and Huitu O. (in press). Non-linear effects of climate on boreal rodent dynamics: mild winters do not negate high-amplitude cycles. Global Change Biology.

Kos M., Hoetmer A. J., Pretorius Y., Boer W. F., de Knegt H. J., Grant C. C., Kohi E., Page B., Peel M., Slotow R., Waal C., Wieren S. E., Prins H. H. T., and Langevelde F. (2012). Seasonal diet changes in elephant and impala in mopane woodland. Eur. J. Wildl. Res. 58:279-287.

Koskinen J. P. and Holm L. (2012). SANS: high-throughput retrieval of protein sequences allowing 50% mismatches. Bioinformatics. 28:i438-i443.

Koskinen P., Laine P., Niemi O. A., Nykyri J., Harjunpää H., Auvinen P., Paulin L., Pirhonen M., Palva E. T., and

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Holm L. (2012). Genome Sequence of Pectobacterium sp. Strain SCC3193. J. Bacteriol. 194:6004.

Kosonen L., Schigel D., Setälä R., and Smirnoff I. (2012). Sieni pusertaa puusta lumihiuksia. Suomen luonto. 71:10-10.

Kraft T. S. and van Nouhuys S. (in press). The effect of host density and species on superparasitism and sex ratio in a gregarious parasitoid. Ecol. Entomol.

Kujala H. (2012). Climate change, species range shifts and uncertainty - A new era of conservation planning. Univerity of Helsinki, Unigrafia, Helsinki.

Kujala H., Burgman M. A., and Moilanen A. (2013). Treatment of uncertainty in conservation under climate change. Conservation Letters. doi: 10.1111/j.1755-1263X.2012.00299.x.

Kujala H., Moilanen A., Araújo M. B., and Cabeza M. (in press). Conservation planning with uncertain climate change projections. PLoS ONE.

Kujala H., Vepsäläinen, V., Zuckerberg, B., and Brommer, J.E. (2013). Range margin shifts of birds revisited – the role of spatiotemporally varying survey effort. Global Change Biology. 19:420-430.

Kukkala A. and Moilanen A. (in press). The core concepts of spatial prioritization in systematic conservation planning. Biological Reviews. doi: 10.1111/brv.12008.

Kurhinen J., Kulebyakina E., Zadiraka E., Mamontov V., Muravskaya E., and Hanski I. K. (2012). Distribution of siberian flying squirrel (Pteromys volans l.) in taiga isthmuses between Baltic and White sea regions. Acta Zool. Litu. 21:306-310.

Kvist J., Wheat C.W., Kallioniemi E., Saastamoinen M., I. H., and M. F. (2013). Temperature treatments during larval development reveal extensive heritable and plastic variation in gene expression and life history traits. Mol. Ecol. 22:602-619.

Laitila J. and Moilanen A. (2012). Use of many low-level conservation targets reduces high-level conservation performance. Ecol. Model. 247:40-47.

Laitila J., Nieminen P. J., Saksman E., and Tylli H.-O. (2013). Compact and weakly compact composition operators on BMOA. Complex Analysis and Operator Theory. 7(1):163-181.

Laurentz M., Reudler J. H., Mappes J., Friman V., Ikonen S., and Lindstedt C. (2012). Diet Quality Can Play a Critical Role in Defence Efficacy Against Parasitoids and Pathogens in the Glanville Fritillary (Melitaea Cinxia). J. Chem. Ecol. 38:116-125.

Leathwick J. R., West D. W., and Moilanen A. (2012). Development of a systematic, information-based approach to the identification of high value sites for freshwater conservation in New Zealand. Pages 183-192 in P. J. B. a. P. J. Raven, editor. River Conservation and Management. Wiley-Blackwell

Liu X., Suzuki A., and Schigel D. (2012). Editorial: The impact of fungi on other organisms. Mycology. 3:1.

Maron M., Hobbs R. J., Moilanen A., Matthews J. W., Christie K., Gardner T. A., Keith D. A., Lindenmayer D. B., and McAlpine C. A. (2012). Faustian bargains? Restoration realities in the context of biodiversity offset

policies. Biol. Conserv. 155:141-148.Martinez J., Duplouy A. M. R., Woolfit M., Vavre F., O’Neill

S. L., and Varaldi J. (2012). Influence of the Virus LbFV and of Wolbachia in a Host-Parasitoid Interaction. PLoS ONE. 7(4).

Mattila A., Duplouy A. M. R., Kirjokangas M., Lehtonen R. J., Rastas P., and Hanski I. (2012). High genetic load in an old isolated butterfly population. Proc. Natl. Acad. Sci. U. S. A. 109:E2496-2505.

Meller L., Barbet-Massin M., Deetman S., Hof A., Pironon S., Thuiller W., and Cabeza M. (2012). Joint impacts of climate change and bioenergy production on the state and conservation of European birds. RESPONSES Project deliverable: Integrated Activity Report I. part of D 5.3.

Mikkonen N. and Moilanen A. (in press). Identification of top priority areas and management landscapes from a national Natura 2000 network. Environmental Science & Policy. 27:11-20.

Miraldo A., Faria C., Hewitt G. M., Paulo O. S., and Emerson B. C. (2012). Genetic analysis of a contact zone between two lineages of the ocellated lizard (Lacerta lepida Daudin 1802) in south-eastern Iberia reveal a steep and narrow hybrid zone. J. Zool. Syst. Evol. Res.

Miraldo A., Hewitt G. M., Dear P. H., Paulo O. S., and Emerson B. C. (2012). Numts help to reconstruct the demographic history of the ocellated lizard (Lacerta lepida) in a secondary contact zone. Mol. Ecol. 21:1005-1018.

Moilanen A. (2012). Reserve selection and conservation prioritization. Pages 617-624 in L. Gross, editor. Encyclo-pedia of theoretical ecology. University of California Press.

Moilanen A. (2012). Spatial Conservation Prioritization in Data-Poor Areas of the World. Natureza & Conservacao. 10:12-19.

Moilanen A. (2013). Planning impact avoidance and biodiversity offsetting using software for spatial conservation prioritization. Wildl. Res.: http://dx.doi.org/10.1071/WR12083.

Moilanen A., Anderson B. J., Arponen A., Montesino Pouzols F., and Thomas C. D. (2013). Edge artefacts and lost performance in national versus continental conservation priority areas. Divers. Distrib. 19:171-183.

Moilanen A., Leathwick J. R., and West D. W. (2012). Development of a systematic, information-based approach to the identification of high value sites for freshwater conservation in New Zealand. in P. J. Boon and P. J. Raven, editors. River Conservation and Management. Wiley-Blackwell.

Moilanen A., Meller L., Leppänen J., Montesino Pouzols F., Kujala H., and Arponen A. (2012). Zonation spatial conservation planning framework and software V3.1, User manual.

Mwakiwa E., de Boer W. F., Hearne J. W., Slotow R., van Langevelde F., Peel M., Grant C. C., Pretorius Y., Stigter J. D., Skidmore A. K., Heitkönig I. M. A., de Knegt H. J., Kohi E. M., Knox N., and Prins H. H. T. (2013). Optimization of wildlife management in a large game reserve through waterpoints manipulation: A bio-economic analysis. J. Environ. Manag. 114:352-361.

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Nemri A., Barrett L. G., Laine A. L., Burdon J. J., and Thrall P. H. (2012). Population Processes at Multiple Spatial Scales Maintain Diversity and Adaptation in the Linum marginale - Melampsora lini Association. Plos One. 7.

Nieberding C. M., Fischer K., Saastamoinen M., Allen C. E., Wallin E. A., Hedenstrom E., and Brakefield P. M. (2012). Cracking the olfactory code of a butterfly: the scent of ageing. Ecol. Lett. 15:415-424.

Niitepõld K. and Hanski I. (2013). A long life in the fast lane: positive association between peak metabolic rate and lifespan in a butterfly. J. Exp. Biol.: doi:10.1242/jeb.080739.

Nilsson R. H., Tedersoo L., Abarenkov K., Ryberg M., Kristiansson E., Hartmann M., Schoch C. L., Nylander J. A. A., Bergsten J., Porter T. M., Jumpponen A., Vasihampayan P., Ovaskainen O., Hallenberg N., Bengtsson J., Eriksson M., Larsson K. H., Larsson E., and Kõljalg U. (2012). Five simple guidelines for establishing basic authenticity and reliability of newly generated fungal ITS sequences. MycoKeys. 4:37-63.

Norros V., Penttilä R., Suominen M., and Ovaskainen O. (2012). Dispersal may limit the occurrence of specialist wood decay fungi already at small spatial scales. Oikos. 121:961-974.

Nykyri J., Niemi O. A., Koskinen P., Nokso-Koivisto J. P., Pasanen M., Broberg E. M., Pljusnin I., Törönen P., Holm L., Pirhonen M., and Palva E. T. (2012). Revised Phylogeny and Novel Horizontally Acquired Virulence Determinants of the Model Soft Rot Phytopathogen Pectobacterium wasabiae SCC3193. PLoS Pathogens. 8:e1003013.

Ohdachi S. D., Yoshizawa K., Hanski I., Kawai K., Dokuchaev N. E., Sheftel B. I., Abramov A. V., Moroldoev I., and Kawahara A. (2012). Intraspecific phylogeny and nucleotide diversity of the least shrews, the Sorex minutissimus-S. yukonicus complex, based on nucleotide sequences of the mitochondrial cytochrome b gene and the control region. Mamm. Study. 37:281-297.

Ovaskainen O. (2012). Pelastetaan vanhat aineistot! Luonnon Tutkija. 3:63.

Ovaskainen O. (2012). Strategies for Improving Biodiversity Conservation in the Netherlands: Enlarging Conservation Areas vs. Constructing Ecological Corridors: An expert report ordered by the Dutch Council for the Environment and Infrastructure. An expert report ordered by the Dutch Council for the Environment and Infrastructure.

Penteriani V. and Delgado M. d. M. (2012). There is a limbo under the moon: what social interactions tell us about the floaters’ underworld. Behav. Ecol. Sociobiol.

Pietola L. H., Jero J., Jalkanen R., Kinnari T., Jero O., Frilander M., Pajusola K., Salminen M., and Aarnisalo A. (2012). Effects of p27Kip1- and p53-shRNAs on kanamycin damaged mouse cochlea. World journal of otorhinolaryngology. 2:1-7.

Pouzols F. M., Barriga Barros A., R. Lopez D., and Sánchez-Solano S. (2012). Enabling Fuzzy Technologies in High Performance Networking via an Open FPGA-Based Development Platform. Applied Soft Computing.

12:1440-1450.Pouzols F. M., Burgman M. A., and Moilanen A. (2012).

Methods for allocation of habitat management, maintenance, restoration and offsetting, when conservation actions have uncertain consequences. Biol. Conserv. 153:41-50.

Pouzols F. M. and Lendasse A. (2012). Adaptive kernel smoothing regression for spatio-temporal environmental datasets. Neurocomputing. 90:59-65.

Pretorius Y., Stigter J. D., de Boer W. F., van Wieren S. E., de Jong C. B., de Knegt H. J., Grant C. C., Heitkönig I., Knox N., Kohi E., Mwakiwa E., Peel M. J. S., Skidmore A. K., Slotow R., van der Waal C., van Langevelde F., and Prins H. H. T. (2012). Diet selection of African elephant over time shows changing optimization currency. Oikos. 121:2110-2120.

Radivojac P., Koskinen P., Nokso-Koivisto J., Holm L. et al. (in press). A large scale evaluation of computational protein function prediction. Nat. Meth.

Saarinen S., Vahteristo P., Lehtonen R., Aittomaki K., Launonen V., Kiviluoto T., and Aaltonen L. A. (2012). Analysis of a Finnish family confirms RHBDF2 mutations as the underlying factor in tylosis with esophageal cancer. Familial Cancer. 11:525-528.

Saastamoinen M., Hirai, N., van Nouhuys, S. (2013). Direct and trans-generational responses to food deprivation during development in the Glanville fritillary butterfly. Oecologia. 171:93-104.

Saastamoinen M., Brakefield P. M., and Ovaskainen O. (2012). Environmentally induced dispersal-related life-history syndrome in the tropical butterfly, Bicyclus anynana. J. Evol. Biol. 25:2264-2275.

Saastamoinen M., Ikonen S., Wong S. C., Lehtonen R., and Hanski I. (in press). Plastic larval development in a butterfly has complex environmental and genetic causes and consequences for population dynamics. J. Anim. Ecol.

Schigel D. (2012). Fungivory and host associations of Coleoptera: a bibliography and review of research approaches. Mycology: An International Journal on Fungal Biology. 3:258-272.

Schigel D. (2012). The impact of fungi on other organisms. Mycology: An International Journal on Fungal Biology. 3:1-1.

Schigel D. S. (2012). Fungivory of saproxylic Coleoptera: the mystery of rejected polypores. In: Saproxylic beetles in Europe: monitoring, biology and conservation. Studia forestalia Slovenica. 137:53–58.

Sharafia S. M., Moilanen A., White M., and Burgman M. (2012). Integrating environmental gap analysis with spatial conservation prioritization: A case study from Victoria, Australia. J. Environ. Manag. 112:240-251.

Sirkiä S. M., Lehtomäki J., Lindén H., Tomppo E., and Moilanen A. (2012). Spatial conservation prioritization of capercaillie (Tetrao urogallus) lekking landscapes in South-Central Finland. Wildl. Biol. 18:337-353.

Sundell J., Church C., and Ovaskainen O. (2012). Spatio-temporal patterns of habitat use in voles and shrews modified by density, season and predators. J. Anim. Ecol.

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Synopsis of the year

81:747-755.Susi H. (2012). Mustavadelman nekroosiviruksen

tunnistaminen. Kasvinsuojelulehti. 3/2012.Taberlet P., Zimmermann N. E., Englisch T., Tribsch A.,

Holderegger R., Alvarez N., Niklfeld H., Coldea G., Mirek Z., Moilanen A., Ahlmer W., Marsan P. A., Bona E., Bovio M., Choler P., Cieslak E., Colli L., Cristea V., Dalmas J. P., Frajman B., Garraud L., Gaudeul M., Gielly L., Gutermann W., Jogan N., Kagalo A. A., Korbecka G., Kupfer P., Lequette B., Letz D. R., Manel S., Mansion G., Marhold K., Martini F., Negrini R., Nino F., Paun O., Pellecchia M., Perico G., Piekos-Mirkowa H., Prosser F., Puscas M., Ronikier M., Scheuerer M., Schneeweiss G. M., Schonswetter P., Schratt-Ehrendorfer L., Schupfer F., Selvaggi A., Steinmann K., Thiel-Egenter C., van Loo M., Winkler M., Wohlgemuth T., Wraber T., Gugerli F., and Consortium I. (2012). Genetic diversity in widespread species is not congruent with species richness in alpine plant communities. Ecol. Lett. 15:1439-1448.

Tack A. J. M., Gripenberg S., and Roslin T. (2012). Cross-kingdom interactions matter: fungal-mediated interactions structure an insect community on oak. Ecol. Lett. 15:177-185.

Tack A. J. M., Johnson M. T. J., and Roslin T. (2012). Sizing up community genetics: it’s a matter of scale. Oikos. 121:481-488.

Tack A. J. M., Thrall P. H., Barrett L. G., Burdon J. J., and Laine A. L. (2012). Variation in infectivity and aggressiveness in space and time in wild host-pathogen systems: causes and consequences. J. Evol. Biol. 25:1918-1936.

Thomas C. D., Anderson B. J., Moilanen A., Eigenbrod F., Heinemeyer A., Quaife T., Roy D. B., Gillings S., Armsworth P. R., and Gaston K. J. (in press). Reconciling biodiversity and carbon conservation. Ecol. Lett.: DOI: 10.1111/ele.12054.

Thrall P. H., Laine A. L., Ravensdale M., Nemri A., Dodds P. N., Barrett L. G., and Burdon J. J. (2012). Rapid genetic change underpins antagonistic coevolution in a natural host-pathogen metapopulation. Ecol. Lett. 15:425-435.

Tollenaere C., Susi H., Nokso-Koivisto J., Koskinen P., A. Tack P. A., Paulin L., Frilander M. J., Lehtonen R., and Laine A.-L. (2012). SNP Design from 454 Sequencing of Podosphaera plantaginis Transcriptome Reveals a Genetically Diverse Pathogen Metapopulation with High Levels of Mixed-Genotype Infection. PLoS One. 7:e52492.

Travis J., Mustin K., Barton K., Benton T., Clobert J., Delgado M. d. M., Dytham C., Hovestadt T., Palmer S., van Dyck H., and Bonte D. (2012). Modelling dispersal: an eco-evolutionary framework incorporating emigration, movement, settlement behaviour and the multiple costs involved. Methods in Ecology and Evolution.

Triviño M. (2012). Global change impacts and conservation priorities in the Iberian Peninsula. University Rey Juan Carlos and National Museum of Natural Sciences, Madrid.

van den Heuvel J., Saastamoinen M., Zwaan B. J., Brakefield P. M., Kirkwood T., and Shanley D. (in press). The

predictive adaptive response: modeling the life history of a tropical butterfly (Bicyclus anynana). Am. Nat.

van Nouhuys S. (2012). Parasiternas fascinerande värld. exempel från åländska ängsmarker.22-24.

van Nouhuys S., Niemikapee S., Hanski, I. (2012). Variation in a host-parasitoid interaction across independent populations. Insects. 3:1236-1256.

van Nouhuys S. and Kraft T. S. (2012). Indirect interaction between butterfly species mediated by a shared pupal parasitoid. Popul. Ecol. 54:251-260.

van Nouhuys S., Reudler J. H., Biere A., and Harvey J. A. (2012). Performance of secondary parasitoids on chemically defended and undefended hosts. Basic Appl. Ecol. 13:241-249.

Van Teeffelen A., Lung T., Meller L., Vermaat J., Lavalle C., and Cabeza M. (2012). EU Biodiversity policy in times of climate change. RESPONSES Policy Update. No 1.

Van Teeffelen A. J. A., Vos C. C., and Opdam P. (2012). Species in a dynamic world: Consequences of habitat network dynamics on conservation planning. Biol. Conserv. 153:239-253.

Virkkala R., Heikkinen, R.K., Fronzek, S., Kujala, H., and Leikola, N. (2013). Does the protected area network preserve bird species of conservation concern in a rapidly changing climate? Biodivers. Conserv. 22:459-482.

Honours and awardsAnna-Liisa Laine received L’Oreal UNESCO for Women

in Science award and Academy of Finland award for Scientific Courage.

VisitorsDavid Muru, February - September 2012.Tanya Semenova, August 2011 - February 2012.Dmitry Finkelstein , January 22 - 28.Dan Lawson, January 22 - 24.Cristina Banks-Leite, February 22 - 23.Robert Ewers, February 22 - 23.Patrik Nosil, February 29 - March 1.Greta Bocedi, March 25 - April 5.Eliezer Gurarie, April 17 - May 4.John Drake, May 22 - 26.Steve Beissinger, April 26 - May 31.Brian Huntley, May 30 - June 3.Felix Horns, August 2011 - June 2012.Benoît Pernechele, February - JuneViacheslav Spirin, April - June.Mamy Rakotoarijaona, August 24 - September 3Juan M. Morales, September 3 - 24Evgen Dykyi, Oleksandr Ordynets, Mariia Pavlovska and

Kseniia Tuholukova, October 7 - 10.

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MRG -ANNUAL REPORT 201250

Statistics

Biostatistiikka II, University of Helsinki, November - December

A basic course in statistics.

Organizer: Otso Ovaskainen

Workshops and meetings

European boreal forest biodiversity (EBFB)-Workshop on Collecting and Analyzing Winter Track Data, Mekrijärvi Biological Station 11. - 13.4.2012

Members of organizing committee: Juri Kurhinen and Otso Ovaskainen

European boreal forest biodiversity (EBFB) - PhD course in St Petersburg on Mathematical and Statistical Modeling of Ecological Data 23. – 27.4.2012

Organizers: Andrei Brodsky and Otso Ovaskainen

Stochastic Dynamics: Mathematical Theory and Applications reserarch program, Bielefeld, Germany, 18.5. - 14.7.2012

Workshops “Stochastic Dynamics in Action” (20–25 May 2012) and “Qualitative behaviour of stochastic systems and applications” (18th June - 21th June)

Member of organizing committee: Otso Ovaskainen

Conservation biology

Reserve planning in the tropics: RESPECT, Madagascar, November - December

This is a CIMO North-South-South initiative. A field course including Malagasy and Finnish st udents, aimed at teaching both ecological and development needs in conservation planning.

Coordinators: Mar Cabeza and Jukka T. Lehtonen

TeachingEcology

Acroecology, Department of Agricultural Sciences, University of Helsinki

Guest lecture on pathogen epidemiology and evolution in the agro-ecological interface

Lecturer: Ayco Tack

Spring Symposium 2012, Departments of Biosciences and Environmental Sciences, Faculty of Agriculture and Forestry, Museum of Natural History

Annual event organised by and for graduate students working in various fields of ecology, evolution, systematics and nature conservation.

Member of organizing committee: Tanjona Ramiadantsoa

Ecology and Evolutionary biology weekly Seminar Series

Organizer: Anna-Liisa Laine & Heikki Hirvonen

Plant-microbe interactions and molecular defence of plants

An advanced course in molecular plant pathology.

Lecturer: Anna-Liisa Laine

Genetics

Next Generation Genomics, University of Helsinki, 6 - 14 September

Two practical courses (mapping and assembly of next generation sequencing data; RNAseq analysis using R) and scientific symposium on next generation sequencing methods.

Member of organizing committee: Mikko Frilander

mRNA processing in eukaryotes, University of Helsinki 12 March - 3 May

Eukaryotic gene expression pathway from the RNA processing point of view. A 3 ECTS undergraduate course.

Teacher: Mikko Frilander

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Synopsis of the year

Conservation Biology in Fragmented Landscapes, University of Helsinki, September - November

Since the year 2000, the Metapopulation Research Group has annually organized a master level course called Conservation Biology in Fragmented Landscapes (7 ECTS). The course is part of the international Boreal Biota and Ecology program opened to both international exchange students and Finnish students. Our objective is to unite all MRG members in a joint venture, to compile a coherent teaching unit that takes advantage of our own research, and to provide group members with essential teaching experience. As in all our work, we blend theory and modelling with empirical case studies, and try to offer the students our view on how habitat fragmentation affects key aspects of population biology. Students are also given the choice of earning two extra credits by reading books produced in the group - this year The Shrinking World by Ilkka Hanski.

In 2012, the course was coordinated by Anne Duplouy, Johanna Eklund and Anniina Mattila. This year we had 15 Finnish students and 12 international students representing ten different nationalities. Following an initiative launched several years ago, we supplement traditional lectures with a series of discussion groups leading up to an informal seminar. These discussion groups, mentored by members of our group, challenge students to apply information learned during the lectures to real world conservation issues. Students are encouraged to evaluate the relevance and scope of existing scientific information from the perspective of applied management issues including the

extent to which scientific information is actually used when real decisions are made. The students presented the results of these working groups during a two day seminar in November at the Lammi Biological Station. To stimulate discussion and introduce a more applied perspective, we invited three external guests working on applied conservation issues: Mia Valtonen from the University of Eastern Finland who is an expert on

the Saimaa seal, Lauri Kajander, who works with forest conservation issues in Luonto-Liitto (the Finnish Nature League), and Juha Lumme, who is an expert on the flying squirrel and works for Uudenmaan ELY-keskus (the Center for economic development, transport

and environment). During the seminar the students gave two presentations per group (a detailed talk aimed at a scientific audience and a shorter one aimed at a more general audience varying from land owners to policy makers). The seminar was a great success, with participants engaging in vigorous discussion and participating in the real-life metapopulation game and other (pseudo)-scientific and purpose-invented games. We also try to implement and improve the course after each year, according to the feedbacks received from the students and the teaching staff. Overall, we find it an exciting challenge to use our own research results to illustrate fundamental ecological concepts to the students. Our teaching unit also functions as a uniting factor, giving us the chance to relate all the work conducted in our group to a unified framework.

Coordinators: Anne Duplouy, Johanna Eklund and Anniina Mattila

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MRG -ANNUAL REPORT 201252

The annual meeting was held 3-7 October in St Petersburg, Russia. We travelled by boat to enjoy the views on the archipelago and to take advantage of the conference facilities, thus starting the meeting already while traveling. The program of the annual meeting consisted of scientific presentations covering both past and planned work, as well as of discussions on how to make our group function even better than it has done so far, both in terms of scientific and non-scientific activities. The regular annual meeting was combined with a Scientific Advisory Board (SAB) meeting that was related to the Centre of Excellence funding from the Academy of Finland. Thus we had on

Annual meeting in St Petersburg

Everybody agrees with Rainer! Yes, let’s go for it!

On a visit to the State Hermitage Museum

board also our SAB members, prof. Marian Goldsmith, prof. Tony Ives and prof. Kevin Gaston, as well as the Academy of Finland representative Harri Hautala. We wish to thank them all once again for fruitful discussions and ideas!

Concerning the social activities, we enjoyed splendid meals both in Georgian and Russian restaurants. As evidenced by the photos, some of us celebrated our scientific achievements all night long, while others focused more on cultural activities.

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Synopsis of the year

Funding

Funding source EUR

AcAdemy of finlAnd

CEnTRE oF ExCELLEnCE FunDInG Metapopulation Research Group 1 077 974

GEnERAL RESEARCH GRAnTSIlkka Hanski: The Glanville Fritillary Butterfly Genome and Population Genomics 116 190

Otso Ovaskainen: Linking Environmental Change to Biodiversity Change: Long-term and Large-scale Data on European Boreal Forest Biodiversity

156 236

ACADEMy PRoFESSoR'S RESEARCH CoSTSIlkka Hanski: Eco-evolutionary spatial dynamics 281 072

ACADEMy RESEARCH FELLow'S RESEARCH CoSTSAnna-Liisa Laine: Studying pathogen evolution: from molecules to metapopulations 133 600

Saskya van Nouhuys: Host-parasite Metacommunity Biology 64 017

Christopher Wheat: Ecological and Evolutionary Functional Genomic Study of Metapopulation Dynamics

81 987

Mar Cabeza Jaimejuan: Conservation efectiveness: Sociopolitical and Environmental challenges, Academy of Finland

2 776

RESEARCH InFRASTRuCTuRES (FIRI 2010)Ilkka Hanski: FIRI:GenoEvo 199 500

PoSTDoCToRAL PRoJECTSAnni Arponen: Coping with Incomplete Data in Conservation Assessments and Policy Tools 91 236

Maria Delgado: Linking Dispersal Strategies with Population Dynamics 96 840

Marjo Saastamoinen: Environmental Stress and Its Effects on Life History Evolution in Wild Populations

104 690

Dmitry Schigel: Colonization gates and establishment of wood-decaying fungi in European Spruce

34 041

+ 1 Academy professor post and 4 Academy research fellow posts

University of HelsinkiCentre of Excellence funding from Ministry of Education 508 933

Ilkka Hanski: Biocentrum Helsinki 46 000

Atte Moilanen: Quantitative Methods for Targeting Conservation Management andIntensive Forestry in Forests in Finland (three year grant)

42 000

Atte Moilanen: ERC/ Moilanen (Department of Biosciences) 45 000

Atte Moilanen: ERC/ Moilanen (Bursar) 25 000

Anna-Liisa Laine: ERC/ Laine (Bursar) 25 000

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MRG -ANNUAL REPORT 201254

Otso Ovaskainen: Starting Grant of the University of Helsinki Funds for Appointment as Professor of Mathematical Ecology

5 000

Atte Moilanen: Starting grant of the University of Helsinki Funds for Appointment as Professor of Conservation Decision Analysis

6 250

Ilkka Hanski: FIRI:GenoEvo (Bursar) 85 500

Swee Chong Wong: GPBM Helsinki Graduate Program in Biotechnology and Molecular Biology, International PhD Training Program

13 770

Henna Fabritius: the University of Helsinki Funds Professors Wives 1 500

+ 2 Laboratory technician posts and 1 professor posts

eUropeAn Union

EuRoPEAn RESEARCH CounCIL

Ilkka Hanski: Ecological, Molecular and Evolutionary Spatial Dynamics 567 600

Otso Ovaskainen: Spatial Ecology: Bringing mathematical theory and data together 300 200

Atte Moilanen: Global Environment Decision Analysis 300 000

Anna-Liisa Laine: Linking Pathogen Evolution and Epidemiology 211 100

CoLLABoRATIVE PRoJECT

Atte Moilanen: Securing the Conservation of Biodiversity Across Administrative Levels and Spatial, Temporal and Ecological Scales

10 000

Mar Cabeza: European Responses to Climate Change: Deep Emissions, Reductions and Mainstreaming of Mitigation and Adaptation

212 141

otHer

FICS - FInnISH DoCToRAL PRoGRAMME In CoMPuTATIonAL SCIEnCESOtso Ovaskainen: Graduate school funding Sonja Koskela 2 486

EMIL AALTonEn FounDATIonHenna Fabritius: Support grant 5 000

FInnISH FoREST AnD PARK SERVICEAtte Moilanen: Support for Professorship in Conservation Decision Analysis. GIS-based Ecological Decision Analysis in the South-Central Finland Forest Biodiversity Program (METSO)

60 000

FInnISH nATuRE ConSERVATIon FounDATIonHenna Fabritius: Succession of Melitaea diamina meadows 1 500

nERC unIVERSITy oF EAST AnGLIAOtso Ovaskainen: Subcontract Agreement: The role of Dispersal in Species Ability to Respond to Climate Change

7 021

nwo nETHERLAnDS oRGAnISATIon FoR SCIEnTIFIC RESEARCHHenjo de Knegt: Rubicon grant 70 328

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Synopsis of the year

Other3 %

Sources of Metapopulation Research Group funding in 2012.

oSKAR ÖFLunD FounDATIonSilvija Budaviciute: Support grant 6 000

SoCIETAS EnToMoLoGICA HELSInGFoRSIEnSISSilvija Budaviciute: Support grant 1 000

THE CEnTRES FoR EConoMIC DEVELoPMEnT, TRAnSPoRT AnD EnVIRonMEnT

Henna Fabritius: Research Collaboration with South Ostrobothnia Centre, Monitoring of the Distribution of Melitaea diamina in Western Finland

3 000

Henna Fabritius: Research Collaboration with Southwest Finland Centre, Monitoring of the Distribution of Melitaea diamina in Western Finland

2 000

LuoVA-GRADuATE SCHooL5 PhD student posts

totAl 5 006 488

All grants from Academy of Finland and EU and some of other funding include overheads. University of Helsinki funding does not include overhead.

European Union

32 %

University of Helsinki

16 %

Academy of Finland

49 %

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MRG -ANNUAL REPORT 201256

Our new funding cycle as a national Centre-of-Excellence (CoE) started in January 2012, and this support will continue until

2016. That is the good news. The bad news is that the level of funding is disappointing. The support from our host institute, the University of Helsinki, is at the same level as in the beginning of our previous CoE period, in 2006, in spite of MRG now being roughly twice as large, and in spite of us bringing roughly 2 million € more overheads per 6 years than in 2006. And this is not all. Although the section of Ecology and Evolutionary Biology (EEB) at our department is performing really well in research and teaching, to a large extent thanks to MRG, the funding situation has deteriorated severely also at this level. The professors who have left, and those whose salary is paid by the research council, are not replaced, which means that the more successful EEB and MRG are, the more is

teaching of ecology and evolutionary biology suffering. Times are tough in the university, but it is inexcusable that the ‘reward’ of success in raising external funding has a negative sign, leads to reduction of core funding.

Putting aside these very troubling prospects concerning university funding for the year 2013, the prospects for MRG are generally good. Several PhD students will finish their studies in 2013, compensating for the past two years with exceptionally low numbers, partly reflecting the shift towards us recruiting more post docs rather than students a few years ago (see the graph on p. 6). We didn’t manage to publish the genome of the Glanville fritillary in 2012 as expected – but surely this will happen in 2013! I personally look forward to the next MRG dung beetle expedition, this time to Gunung Mulu in Sarawak, Malaysia, where I did field work back in 1978. It is very special to go back and to have a chance to take some of you with me.

Ilkka Hanski

Prospects for the year 2013