Annual Report 2012

Metapopulation Research Group

Department of Biosciences Faculty of Biological and Environmental Sciences University of Helsinki

Helsinki 2012 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: www.helsinki.fi/science/metapop

Layout: Sami Ojanen MRG-logo: Gergely Várkonyi Face photos: Evgeniy Meyke / Sami Ojanen

Printed in Picaset Oy

Helsinki, January 2013 Contents

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

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

& Research Projects...... 9 • The Glanville fritillary model system...... 10 Metapopulation biology of the Glanville fritillary butterfly: Ecological and evolutionary spatial dynamics 10 Genomics and genetics of the Glanville fritillary butterfly 12 Environmental stress and its effects on life history evolution in wild populations 16

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

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

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

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

& Supporting personnel...... 38

& Synopsis of the year 2012...... 41 Publications 42 Honours and awards 49 Visitors 49 Teaching 50 Annual meeting in St Petersburg 52 Budget 53 & Prospects for the year 2013...... 52

Foreword

Happy researchers

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

5 Brief history and overview of MRG

etapopulation Research Group was plant-mildew system studied by Anna-Liisa Laine, the established by Ilkka Hanski 21 years ago, in community of wood-decomposing fungi studied by 1991. Ilkka had worked on the ecology of Otso Ovaskainen, and the radiation of endemic dung spatiallyM structured populations since the late 1970’s. beetles in Madagascar studied by Ilkka. Atte Moilanen’s The early work was concerned with small-scale spatial Zonation software has attained worldwide reputation aggregation of individuals within populations and how in systematic conservation planning. In the past, MRG that might affect the coexistence of competitors. Since was primarily a research group in ecology, but this is the early 1980’s the focus had shifted to larger spatial no longer the case. The senior researchers have opened scales and to the dynamics of metapopulations, networks up new fronts of research to bring together population of local populations with relatively independent biology and state-of-the-art mathematical modeling demographic dynamics. Significant events leading to the (Otso Ovaskainen), molecular biology (Mikko establishment of MRG included the first international Frilander) and genomics (Rainer Lehtonen), as well workshop on metapopulation ecology organized by as developed new approaches to conservation biology Ilkka and Michael Gilpin (San Diego, US) in 1989, and reserve design (Atte Moilanen, Mar Cabeza). The which resulted in the first edited volume on the subject metapopulation concept is no longer as fundamental to (Gilpin & Hanski, 1991, Metapopulation Dynamics: our research than it was in the past, though processes Empirical and Theoretical Investigations, Academic related to the spatial structure and dynamics of Press, London). The long-term metapopulation study of populations remain the focus of much of our research. the Glanville fritillary butterfly in the Åland Islands in Finland was started in 1991. Current structure

Current research The graph shows the growth of MRG since 1992. At present, MRG is a highly international group of 73 MRG is the leading research group worldwide in researchers (9), post docs (28), post graduate students metapopulation biology and one of the Centres-of- (23), and supporting personnel (13) representing 19 Excellence in research nominated by the Academy of different nationalities. MRG currently consists of seven Finland (national research council) for 2000-2005, research groups with their own students and post 2006-11 and again for 2012-2016. Our past strengths docs and largely own funding. We are united by the include successful integration of theory, modeling and shared Centre-of-Excellence funding, which supports empirical studies in the same projects. The Glanville a common office and other facilities. Equally, we are fritillary butterfly has become a widely recognized united by much interaction among the research groups model system in population biology, but other long- around four major research themes: term projects are increasing in prominence: the host- I Local adaptation, ecological and coevolutionary parasitoid system studied by Saskya van Nouhuys, the dynamics, and evolutionary radiations II Genomics, genetics and functional molecular biology III Mathematical ecology IV Ecological decision analysis and applied conservation The 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

Fig. 1. MRG personnel since 1992. The objective is to advance the general understanding of the dynamics of species living in heterogeneous environments, with explicit attention to interactions

6 MRG -ANNUAL REPORT 2012 Overview between demographic dynamics and local adaptation, Second, we aim to relate these theories to empirical between behaviour and ecological dynamics in data via new statistical approaches. Third, many of the metacommunities, and between the dynamics of species’ empirical projects described in Themes I and II will be geographical ranges and evolutionary radiations. Many complemented with more specific modelling projects. projects involve molecular studies and modelling. Fourth, we will develop modelling tools to better connect decision analysis and conservation (Theme IV) to basic population and community ecology. Theme II – Genomics, genetics and functional molecular biology The main objective is to develop and implement Theme IV – Ecological decision analysis and genomic and genetic tools to the study of natural applied conservation populations of non-model species. Most of the work This theme has four objectives, all building on past is focused on the Glanville fritillary, but other study research by the senior researchers in MRG. First, we projects also benefit of the methods developed. The develop methods for conservation prioritization that current main task is to complete the draft genome of the integrate across multiple environments and levels of Glanville fritillary, which will greatly facilitate research biological organization. Second, we consolidate the on this major study system in the future. linkage between conservation planning and cutting- edge species-level and community-level spatial Theme III – Mathematical ecology modelling. Third, we improve the understanding of how climate change should be accounted for in conservation. This theme has four objectives. First, we aim to develop Fourth, we develop methods for the evaluation of general theory in spatial ecology, genetics and evolution. conservation outcomes.

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

Ilkka Hanski Otso Ovaskainen Atte Moilanen Rainer Lehtonen Guillaume Blanchet Enrico Di Minin Virpi Ahola Henjo De Knegt 3 Jussi Laitila Anton Chernenko Maria Delgado Federico Pouzols Maaike De Jong Juri Kurhinen Tuuli Toivonen Anne Duplouy Juho Pennanen Peter Kullberg Andreia Miraldo Dmitry Schigel Joona Lehtomäki 3 Pasi Rastas Daniel Simpson Panu Somervuo Ulisses Camargo Mar Cabeza Patrik Koskinen 1 Jussi Jousimo Anni Arponen Anniina Mattila Markku Karhunen 4 Astrid van Teeffelen Swee Chong Wong Sonja Koskela Silvija Budaviciute Veera Norros Johanna Eklund Anna-Liisa Laine Tanjona Ramiadantsoa 3 Henna Fabritius Benoit Barres Rachel Garcia Kristina Karlsson 2 Saskya van Nouhuys Heini Kujala Hannu Mäkinen Delia Pinto Zevallos Laura Meller Ayco Tack Christelle Couchoux Maria Triviño 5 Charlotte Tollenaere Wolfgang Reschka Laure Zupan Riikka Alanen Hanna Susi Mikko Frilander Chris Wheat Jouni Kvist 3 Emily Hornett Marjo 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,

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

Metacommunity dynamics Metacommunity dynamics of wood-decaying fungi Parasitoid population ecology

Coevolutionary dynamics and radiations Species Interactions in Metapopulations Evolutionary radiations of dung beetles in Madagascar

Mathematical ecology Modelling dispersal and population dynamics European Boreal Forest Biodiversity (EBFB)

Systematic conservation planning Biodiversity conservation informatics Conservation Effectiveness Climate Change

Supporting personnel

Maaike de Jong Ilkka Hanski Anne Duplouy Post doc Project leader Anniina Mattila Post doc PhD -student

he large metapopulation of the Glanville Highlights of the year fritillary butterfly (Melitaea cinxia) in the Åland Islands in southwestern Finland has Anne Duplouy has conducted a common garden study Tbeen studied since 1991. Over the years, the Glanville of the life history ecology and evolution of four regional fritillary has become a widely recognized model system populations of the Glanville fritillary from two highly not only in metapopulation ecology but in population fragmented landscapes (Åland Islands in Finland and biology more generally. The research has made many Uppland in Sweden) and two continuous landscapes contributions to the study of the ecological, genetic and (Saaremaa Island in Estonia and Öland Island in evolutionary consequences of habitat fragmentation, Sweden). As predicted by previous models, butterflies and the empirical studies have stimulated the from the fragmented landscapes have higher dispersal development of new concepts and models. Currently rate (as measured by peak flight metabolic rate) than the ecological studies are coupled with functional butterflies from continuous landscapes. Anniina Mattila genomics research (p. 12). and others have done research on a completely isolated,

A B

PTxPT Cross ÅLxÅL PT Cross ÅL Population/cross Population/cross 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).

10 MRG -ANNUAL REPORT 2012 The Glanville fritillary model system

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.

small and old (at least 76 years) local population of the correlation between Cyp and host plant preference Glanville fritillary on the island of Pikku Tytärsaari across the Åland Islands (Fig. 3). It may well turn out (PT) in the Gulf of Finland. Despite its high genetic that the dynamics of genetic variation in the Cyp gene is load manifested as greatly reduced fitness (Fig. 1), one coupled with ecological dynamics and hence represents striking local adaptation has been documented for the yet another example of eco-evolutionary dynamics in PT population. The claws of the PT butterflies have a the Glanville fritillary. The models of eco-evolutionary greater curvature, apparently improving their grip dynamics that have been stimulated by the Glanville and allowing the butterflies to resist strong air force, fritillary project have been recently applied to another and hence windy conditions on the island, which is study system, the Timema walking stick insects in expected to reduce the risk of being blown off to the sea California, in collaboration with Prof Patrik Nosil. (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 Collaborators heritability for lifetime reproductive success, the key component of fitness. A large part of this heritability Prof Rongjiang Wang, Peking University, China Prof Paul Brakefield, University of Cambridge, UK was explained by a single Cytochrome P450 gene (Cyp), Dr Kristian Niitepõld, Stanford University, USA which is known to be involved in host plant adaptation Prof Patrik Nosil, University of Sheffield, UK in other insect species. The results show a strong 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, Fig. 3. Cyp allele frequency is significantly correlated with 37, E2496-E2505. average host plant preference in the Åland Islands (average Hanski I. (2012). Dispersal and eco-evolutionary dynamics values for different parts of the study area). Here, the in the Glanville fritillary butterfly. In Dispersal Ecology horizontal axis shows the frequency of the G allele in Cyp and Evolution Clobert J., Baguette M., Benton T.G. and and the vertical axis shows preference for Plantago over Bullock J.M. (eds.), pp. 290-303. Oxford University Press, Veronica. Oxford.

11 Genomics and genetics of the Glanville fritillary butterfly

Rainer Lehtonen Mikko Frilander Ilkka Hanski Project leader Senior researcher Senior researcher

Pasi Rastas Panu Somervuo Virpi Ahola Post doc Post doc Post doc

Swee Chong Wong Patrik Koskinen Jouni Kvist PhD -student PhD -student PhD -student

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

he main objective of this research project is to platforms. The current draft genome assembly develop and implement genomic and genetic (table 1, figure 1) consists of 8,262 scaffolds tools for the study of natural populations of (>1,500 bp) and includes approximately 15,000 Tnon-model species. Most of the work is focused on the gene models. Automated annotations have been Glanville fritillary, but other study projects will benefit manually curated for more than 600 genes. of the methods to be developed. We aim at localizing Roche-Nimblegen and SOLiD RAD-tag data from and identifying genes and genetic variants affecting controlled crosses including 22,000 SNPs have phenotypic and life-history traits that influence local been used to construct a genetic map with the adaptation and population dynamics. A population- correct number of chromosomes (n=31). The 1st wide pedigree will be constructed for the Glanville version of the genome will be published in 2013. fritillary using spatial and genetic data. 2. RNA sequencing of population samples and samples from experiments on flight metabolic Highlights of the year rate. We have individually sequenced transcriptomes (RNA) of 198 samples originating 1. We have completed the sequencing of the ~400 from five different regional populations. Åland Mb genome of the Glanville fritillary using Roche and Uppland populations live in a fragmented 454, Illumina GAII/HiSeq2000 and SOLiD4/5500 landscape, whereas Saaremaa and Öland represent

12 MRG -ANNUAL REPORT 2012 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 Influence of library insert size on scaffold N50 size. counted to be correct if they map to one chromosome but Fig. 1. The most significant increase in scaffold size occurred after are assumed to involve assembly errors if mapped to two or the 454 20kb library data (library 9) had been added to the more different chromosomes. assembly.

Collaborators Petri 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 continuous landscapes. Pikku Tytärsaari is a small isolated population suffering from a high genetic Mattila A.L.K., Duplouy A, Kirjokangas M., Lehtonen R., load. An experiment on flight metabolism and Rastas P. and Hanski I. (2012). High genetic load in an differences in flight performance includes 90 RNA old isolated butterfly population. Proc. Natl. Acad. Sci. samples. In-depth analyses of these data sets, 109(37), E2496-505. Kvist J., Wheat C.W., Kallioniemi E., Saastamoinen M., including Gene Ontology (figure 2) and pathway Hanski I. and Frilander M.J. (2013). Temperature enrichment analyses, are on-going. treatments during larval development reveal extensive 3. The genomic information has been used to design heritable and plastic variation in gene expression and life genotyping assays for large-scale association and history traits. Mol Ecol. 22:602-619 linkage studies. Targeted genotyping experiments Saastamoinen M., Ikonen S., Wong S.C., Lehtonen R. have revealed many novel gene-phenotype and Hanski I. Plastic larval development in a butterfly has complex environmental and genetic causes and associations such as a vitellin-degrading protease consequences for population dynamics. J. Anim. Ecol., in precurson gene in larval growth (Ahola et al. in press. prep.) and the occurrence of an extra larval instar Karinen S., Saarinen S., Lehtonen R., Rastas P., Vahteristo (Saastamoinen et al. in press), and a cytochrome P., Aaltonen L.A. and Hautaniemi S. 2012. Rule-based P450 gene affecting the reproductive performance induction method for haplotype comparison and of Glanville fritillary females (de Jong et al. in identification of candidate disease loci. Genome Med. prep.). 4(21).

13 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.

14 MRG -ANNUAL REPORT 2012 The Glanville fritillary model system

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

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, Marjo Saastamoinen however, in both species individuals seem to use Researcher 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 he ability to cope with, react or even adapt and abdomen, which results in increased flight ability to environmental stress is essential for most of the stressed individuals. This response potentially organisms. Yet, surprisingly little is still known allows the stressed individuals to move away from the Tabout the strategies and genetic mechanisms involved in poor habitat patches, as was shown by our theoretical coping with environmental stress in wild populations. model1. In M. cinxia, on the other hand, mothers In this project, I have studied the influence of stressful stressed during development produce offspring that developmental conditions, in terms of thermal and can cope better with similar type of stress during resource levels, on life history variation in two butterfly their own development compared with offspring from species, Bicyclus anynana and Melitaea cinxia. With non-stressed mothers2 (Fig. 1). To compensate for the help of quantitative genetics and candidate gene suboptimal thermal conditions during development, M. approaches I have further aimed to understand cinxia can even increase the number of developmental the (genetic) mechanisms underlying the observed instars3. Variation in instar number seems to be responses. 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).

16 MRG -ANNUAL REPORT 2012 The Glanville fritillary model system

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

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

Collaborators Selected publications

Michelle Baguette, Experimental Ecology Research Centre, 1. van den Heuvel J., Saastamoinen M., et al. (2013). CNRS in Moulis, France The predictive adaptive response: modeling the life Prof. Paul Brakefield, University of Cambridge, UK history of a tropical butterfly (Bicyclus anynana). Jean Clobert, Experimental Ecology Research Centre, American Naturalist, in press. CNRS in Moulis, France 2. Saastamoinen M., Norio H. & Van Nouhuys S. Jon Brommer, University of Turku (2013). Direct and trans-generational responses to Markus Rantala, University of Turku food deprivation during development in the Glanville Prof. Bas Zwaan, University of Wageningen, The fritillary butterfly. Oecologia, in press. Netherlands 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.

17 Metacommunity dynamics of wood-decaying fungi

Otso Ovaskainen Dmitry Schigel Project leader Post doc

Sonja Koskela Veera Norros PhD -student PhD -student Technician: Heini Ali-Kovero

his project is focused on metacommunity dynamics of wood-inhabiting fungi living inT 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 Fig. 1. Woodpecker´s job. To avoid what kind of management measures felling trees in protected areas, would be most effective in improving sampling standing trees requires the viability of wood-inhabiting fungi in ropes and carabiners (left). Samples boreal forests. One key aspect of the work from living trees (above) often contain a mix of hard sawdust is molecular identification of fungi from (light colour) and decayed particles next generation sequencing data, and (brown), suggesting presence of the combination of sequencing data with wood-decaying fungi. sporocarp survey data.

18 MRG -ANNUAL REPORT 2012 Metacommunity dynamics

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 Collaborators

Dmitry obtained a Finnish Academy post doc grant Dr. Üllar Rannik and Dr. Tareq Hussein, University of for the project ”Colonization gates and establishment Helsinki Dr. Panu Halme, University of Jyväskylä of wood-decaying fungi in European Spruce”. This year Dr. Raisa Mäkipää, Dr. Reijo Penttilä, and M.Sc. Juha he sampled living and standing dead trees (Fig 1) to Siitonen, Finnish Forest Research Institute reconstruct the history of fungal arrival by spores. In Prof. Jan Stenlid, M.Sc. Elisabet Eriksson, the Swedish addition to field work in Finland, Dmitry collected University of Agricultural Sciences samples from China, Poland, and Sweden. Supervised Dr. Jenni Nordén and Dr. Karl-Henrik Larsson, University by Dmitry, Virve Koivuranta started her master´s of Oslo project on insects as dispersal vectors of wood- Dr. Henrik Nilsson, University of Gothenburg decaying fungi. At the time of writing this report, Veera is busy Recent publications with finishing her PhD, the deadline for submitting the work to pre-examiners being early January 2013. Liu X., Suzuki A. And Schigel D. (2012) Editorial: The The combination of field measurements, laboratory impact of fungi on other organisms. — Mycology 3: 1. experiments and mathematical modelling has resulted Nilsson R. H., Ovaskainen O., et al. (2012). Five simple in many interesting results. In particular, we have guidelines for establishing basic authenticity and been long interested to learn about the role of spore reliability of newly generated fungal ITS sequences. MycoKeys 4, 37-63. size in fungal dispersal. The collaboration with aerosol Norros V., Penttilä R., Suominen M. and Ovaskainen O. physicists has shown that deposition processes (rather (2012). Dispersal may limit the occurrence of specialist than e.g. gravitational settling) seems to be the key in wood decay fungi already at small spatial scales. Oikos making large spores disperse shorter distances than 121, 961-974. small spores (Fig. 2). Schigel D.S. (2012). Fungivory of saproxylic Coleoptera: the Sonja continued to develop probabilistic methods mystery of rejected polypores. In: Saproxylic beetles in of taxonomic classification necessary for species-level Europe: monitoring, biology and conservation. — Studia identification based on molecular barcoding data - forestalia Slovenica 137: 53–58. such data are now being generated in large quantities Schigel D.S. (2012) Fungivory and host associations thanks to Heini’s work. of Coleoptera: a review of research approaches. — Mycology, 3:258-272.

19 Parasitoid population ecology

Saskya van Nouhuys Delia Pinto Project leader Post doc

Christelle Couchoux Wolfgang Reschka PhD -student PhD -student

PhD -student: Katie Sullivan (Cornell University) Undergraduate students: Aapo Ahola1, Riikka Alanen1, Peter Kullberg1, Terhi Lahtinen1, David Muru, Markus Tietäväinen 1 co-supervised with Anna-Liisa Laine

e study the community, population and aspects of parasitoid (Cotesia melitaearum) virulence, behavioural ecology of parasitoids in a such as brood size, differed among parasitoid populations. spatial context. The project started in the There was however, no evidence of local adaptation Wearly 90s with the spatial dynamics of two parasitoids by either species. There was also no apparent cost of of the Glanville fritillary butterfly in Åland, Finland. inbreeding for the parasitoid from the tiny isolated island Gradually it has broadened to include other parasitoids, of Pikku Tytärsaari. One interesting trait that differed hyperparasitoids, pathogens, symbionts, the host food among populations for both hosts and parasitoids was the plants, and a related butterfly, Melitaea athalia. number of wasp generations per year (Fig. 1), which can have a large impact on their population dynamics. Highlights of the year Previously we have studied the multitrophic effects of toxic chemicals produced by plants on parasitoids. A highlight from the population ecology side of our This year we started working on another aspect of plant research comes from a study of parasitoid virulence chemistry, the volatile organic chemicals (VOCs) that and host susceptibility across five Baltic populations the plants emit. Since herbivores and their parasitoids of the Glanville fritillary butterfly. The ability to resist both use these volatiles as foraging cues, plants are under parasitism differed among butterfly populations, and 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) parasitoid origin cocoons a much different VOC profile than an intact * * plant, and that even oviposition by M. cinxia 3rd instar onto the surface of a leaf causes the volatile emission to change (Fig. 2). The next steps are

1st instar 2nd instar Åland Pikku-Tytärsaari Saaremaa Uppland Fig. 1. The proportion of C. melitaearum in each immature life stage at host diapause stage, b) host origin 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 Åland Öland Saaremaa Uppland at p < 0.05 in the fraction maturing in autumn.

20 MRG -ANNUAL REPORT 2012 Metacommunity dynamics

** 200 *** ** * ** **

150 **

100 Number of caterpillars 50

Fig. 2. Emissions rate of 9 volatile compounds from Veronica spicata that is undamaged (white bars), fed upon (gray 0 bars), and oviposited upon (black bars) by Melitaea cinxia. A B C D E F G H I J Significance between treatments is shown as * P < 0.05; ** P Melitaea cinxia egg cluster in order of size < 0.05; *** P < 0.001. Fig. 4. The number of Melitaea cinxia larvae parasitized by Hyposoter horticola (coloured stripes) and unparasitized to measure the responses of M. cinxia and the parasitoids (gray stripes) in 10 egg clusters parasitized in the field. In to the different volatile profiles. each bar the progeny of each mother wasp is represented by a We have also continued a long-term study of the different colour, so for example all of the parasitoid offspring behavioural ecology of the parasitoid Hyposoter horticola. in cluster F have the same mother whereas in cluster I most The adult females of this species compete strongly, with progeny were mothered by a single wasp, but 7 other mothers multiple females finding and monitoring host egg clusters produced a few progeny each. The parentage of the offspring before they are ready to be parasitized. A new graduate is determined using 14 microsatellite markers. The yellow student, Wolfgang Reschka, is studying how the wasp colour in some of the bars represents wasp progeny that could keeps track of the spatial locations of host egg clusters not be genotyped (due to poor quality DNA extraction), and in a landscape. He will do this by following the foraging therefore could not be assigned to a mother. 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). Collaborators After parasitizing a fraction of the hosts in a cluster Dr. H. Hakola, Finnish Meteorological Institute, FI an individual H. horticola leaves a deterrent mark on the Dr. J. A. Harvey, Netherlands Institute of Ecology, NL plant. This year Christelle Couchoux used microsatellite Prof. J. K. Holopainen, University of Eastern Finland, FI markers to determine the effectiveness of the deterrent. Prof. R. F. Medina, Texas A & M University, USA She found that while one mother wasp parasitizes the Prof. D. Quicke, Imperial College, Silwood Park, UK majority of hosts in a cluster there are, as expected in the Dr. J. H. Reudler, University of Jyväskylä, FI competitive environment, “cheaters,” especially in the Dr. M. S. Shaw, National Museum of Scotland, UK large host clusters (Fig. 4). 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 Fig. 3. The parasitoid wasp Hyposoter horticola with a parasitoids on chemically defended and undefended small piece of metal glued to its thorax. hosts. Basic and Applied Ecology, 13: 241-249

21 Species Interactions in Metapopulations

Anna-Liisa Laine Ayco Tack Project leader Post doc Kristina Karlsson Green * Hanna Susi Post doc PhD -student

Hannu Mäkinen Charlotte Tollenaere Post doc Post doc Benoit Barrès Riikka Alanen Post doc PhD -student Fulbright scholar: Felix Horns Undergraduate students: Aapo Ahola1, Peter Kullberg1, Terhi Lahtinen1, Sini Mursinoff2 , Benoit Pernechele3 * Joint project with Marjo Saastamoinen & Saskya van Nouhuys 1 Supervisors Anna-Liisa Laine & Saskya van Nouhuys 2 Supervisors Anna-Liisa Laine & Ayco Tack 3 Supervisor Charlotte Tollenaere

ur work is focused on uncovering eco- Fig. 1. The number of spores landing on evolutionary links in a large natural microscope slides decreases with distance plant-pathogen metapopulation of from the pathogen source. The black line theO powdery mildew fungus, Podosphaera 10.0 shows the total number of spores and the plantaginis, infecting Plantago lanceolata in the coloured lines show the contribution of different spore group sizes. 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 1.0 in April and has started her PhD work focused on overwintering of Po. plantaginis. During spring

term, Benoit Pernechele (Louvain Academy, Number of spores Belgium) completed his thesis project under the supervision of Charlotte. In May Kristina

Karlsson-Green was awarded the Formas grant 0.1 in Sweden and she joined us to carry out a joint project with Saskya van Nouhuys and Marjo Group size Saastamoinen on the interactions among trophic Total 3−5 levels associated with Pl. lanceolata. During a 1 >5 2 post doctoral project Hannu Mäkinen developed 0 50 100 150 200 250 a qPCR method for detecting and quantifying Distance from source (cm)

22 MRG -ANNUAL REPORT 2012 Coevolutionary dynamics and radiations

●

Time to sporulation (days) 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 8 high variability in the levels of chasmothecia observed among populations. 0 50 100 150 • Hanna’s experimental populations of Pl. lanceolata at Distance from source (cm) the Lammi Biological Station demonstrated that disease Fig. 2. The time to sporulation increases as a transmission is strongly affected by the resistance function of distance from the pathogen source. strategy of the host. Also, we find that infection levels are higher, and decline more rapidly, in populations co-infection. And finally, Benoit Barrès joined us in with co-infection compared to populations infected by October to begin work on molecular epidemiology of a single pathogen strain (Fig. 3). Po. plantaginis. Collaborators: Highlights of the year Dr Jeremy Burdon, CSIRO Plant Industry, Canberra, Australia • This year marked the highest disease prevalence in Prof. David Gadoury, University of Cornell, USA the Åland metapopulation with nearly 700 infected Emilie Haon-Lasportes, INRA, Avignon, France populations. Prof. Levente Kiss, Academy of Sciences, Budapest, • The project on Po. plantaginis transcriptome Hungary sequencing, and development and validation of the Dr Samuel Soubeyrand, INRA, Avignon, France first SNP panel was completed this year through a Dr Peter Thrall, CSIRO Plant Industry, Canberra, Australia successful collaboration of many partners (Tollenaere et al. 2012). We discovered that approximately half of the local pathogen populations contain genetic Key Publications diversity, and in these populations co-infection is Tollenaere C., Susi H., Nokso-Koivisto J., Koskinen P. common. Tack A. J. M., Auvinen P. Paulin L., Frilander M. J., • Ayco’s dispersal experiments using a wind tunnel and Lehtonen R. & Laine A.-L. (2012). SNP Design from 454 common garden setting revealed that each step of the Sequencing of Podosphaera plantaginis Transcriptome transmission process - production of dispersal spores, Reveals a Genetically Diverse Pathogen Metapopulation the distance they travel, and successful colonization with High Levels of Mixed-Genotype Infection. PLoS of a new host – is strongly mediated by interactions ONE, 7:e52492. between host and pathogen genotypes. Interestingly, Tack A., Thrall P. H. Barrett L. G., Burdon J. J. & Laine A.- the size of the dispersal unit varies from individual L. (2012). Variation in infectivity and aggressiveness in spores to spore clumps, and the size of the dispersal space and time in wild host-pathogen systems - causes unit has a striking impact on both dispersal distance and consequences. Journal of Evolutionary Biology 25:1918-1936. (Fig. 1) and speed of development of the new infection Nemri A., Barrett L. G., Laine A.-L., et al. (2012). Mode of (Fig. 2). reproduction predicts population structure at different • Charlotte demonstrated, using both molecular spatial scales in the Linum marginale - Melampsora lini and phenotypic data, that Po. plantaginis is capable association. PLoS ONE 7: e41366. of selfing. The production of chasmothecia was Thrall P. H., Laine A.-L., et al. (2012). Rapid genetic change also affected by ambient temperature and by the underpins antagonistic coevolution in a natural host- interaction between temperature and pathogen pathogen metapopulation. Ecology Letters, 15: 425-35.

23 Evolutionary radiations of dung beetles in Madagascar

Ilkka Hanski Project leader

Andreia Miraldo Tanjona Ramiadantsoa Post doc PhD -student

Undergraduate Kaisa Torppa Student:

here are more than 250 Anton Chernenko sub-clades have expanded to the species of endemic dung Post doc west coast and especially to the east beetles in Madagascar in four coast. The rate of diversification has Tlarge radiations, representing four independent generally declined over time, but the emergence of colonizations during the Cenozoic (one clade may a clade of large-bodied Nanos ca 6 mya is associated date from the late Mesozoic). Since 2002, researchers with a secondary burst of speciation. These species and students in MRG have constructed molecular may have gained competitive advantage because of phylogenies that include most of the described species, their large body size. This lineage has expanded its we have sampled beetles across Madagascar to describe geographical range throughout all of Madagascar their geographical ranges, and we have studied the and speciated into 24 species mostly in allopatry. breeding biology and ecology of selected species and We are analysing transcriptomic data for 9 of the 24 local communities. For a recent review see http://www. species to elucidate the genomic underpinnings of mdpi.com/2075-4450/2/2/112. the radiation. In the past two years, research has been focused Tanjona Ramiadantsoa is developing a stochastic on the radiation consisting of the genera Nanos and occupancy model that combines the extinction- Apotolamprus (Canthoninae). This is the youngest of speciation dynamics (radiation) with the long-term the four major radiations with the estimated age of ca dynamics of species’ geographical ranges. For the 20 my and 74 described species. Andreia Miraldo has purpose of modelling, Madagascar is divided into collaborated with Dr Olivier Montreuil (Paris) and five regions within which extinction, ‘sympatric’ Dr. Heidi Viljanen (Helsinki) to revise the taxonomy speciation and colonization to an unoccupied region of Nanos using both genetic and morphological data. are the possible events. If a lineage occurs in more We are currently analysing the Nanos-Apotolamprus than one region, explicit allopatric speciation is an radiation in detail, examining evolution of body additional possible event. The model will be fitted to sizes, species’ elevational distributions and their the radiation of the Nanos-Apotolamprus clade using geographical ranges. It is apparent that the radiation Approximate Bayesian Computation (ABC) methods. started from North Madagascar, from where many

24 MRG -ANNUAL REPORT 2012 Coevolutionary dynamics and radiations

Nan.bic. Nan.mang Nan.hum. Nan.ate. Nan.con. Nan.min. Nan.psem Ap.cya.. Ap.han.. Ap.quan. Ap.mil.. Ap.vad.. Ap.amb.. Ap.mar.. Ap.lat.. Ap.zom.. Ap.met.. Ap.ser.. Ap.hel.. Ap.hag.. Ap.pseq. Ap.quam. Ap.ora.. Ap.dar.. Ap.per.. Ap.sah.. Ap.pse.. Ap.man.. Nan.sem. Nan.ran. Nan.pun. Nan.pser Nan.rubs Nan.han. Nan.and. Ln (Body-size) : Nan.mar. Nan.pey. 0.92 – 1.17 Nan.pse. A Nan.vad.

1.17 – 1.45 Nan.man. Nanos * increasein diversificationrate Nan.bin. 1.45 – 1.68 Nan.bim. B Nan.occ. 1.68 –1.93 Nan.mag.

Nan.vie. bodied 1.93 – 2.19 Nan.vie4

Nan.nit. ge 2.19 – 2.44 Nan.dub.

Nan.mir. Lar Nan.cly.

Miocene Pliocene Pleistocen.

20.0 17.5 15.0 12.5 10.0 7.5 5.0 2.5 0.0

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.

Collaborators

Olivier Montreuil, the Paris National Museum of Natural History, France Heidi Viljanen, Finnish Natural History Museum, Helsinki Ari 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. Fig. 2. Field work in Madagascar. Preparing fish baits for Miraldo A. and Hanski I. Tempo and mode of trapping dung beetles (which are mostly attracted by carrion diversification in the radiation of Malagasy dung beetles. as well), and a plateful of canthonines, mostly Nanos viettei. In preparation.

25 Modelling dispersal and population dynamics

Henjo De Knegt Guillaume Blanchet Post doc Post doc

Henna Fabritius Maria Delgado PhD -student Post doc

Otso Ovaskainen Project leader

Tanjona Ramiadantsoa Markku Karhunen PhD -student PhD -student

Jussi Jousimo Ulisses Camargo PhD -student PhD -student

athematical modelling can be used in Evolution of measures ecological research in (at least) two ways. First, (model definition, what the individuals do?) mathematical analyses can help to understand Mcausal 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 Evolution of spatial moments this approach to analyze what drives spatial community (how the model behaves at the population level?) 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 Evolution of cumulants on animal movements, with the aim of inferring the (same as above, but a better starting point for analysis) underlying mechanisms from the emergent patterns that can be observed in data.

Highlights of the year Fig. 1. A mathematical framework for analyzing spatio- temporal point processes. Collaboration with mathematicians In 2012 we welcomed Guillaume and Ulisses as new from Bielefeld University (Yuri Kondratiev and his team) group members. Having a background in community- and theoretical ecologists (Stephen Cornell and Ben Bolker) level approaches, Guillaume started to implement into resulted in a mathematical toolbox that applies to a wide range an R-package extensions of community-level models of stochastic and spatial models of interacting individuals.

26 MRG -ANNUAL REPORT 2012 Mathematical ecology

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 that we have developed earlier. Furthermore, in dynamics of the false heath fritillary, to continue with collaboration with Vesa Selonen and others, Guillaume modelling next year. started modelling the broad-scale occurrence of flying Otso spent two months of the summer 2012 in squirrel in Finland. Henjo is also involved in the flying Germany, where he participated (as one of the four squirrel project but he focuses on dispersal from the organizers) the research program “Stochastic Dynamics: point of view of movement analyses. Ulisses started Mathematical Theory and Applications” at ZiF (Center his PhD where his aim is to use a grid of autonomous for Interdisciplinary Research), Bielefeld. These months audio recorders to model the spatio-temporal were especially important to bring mathematical rigor dynamics of birds in the Amazonian rain forests. This to the methods we have developed earlier to analyse project is conducted in collaboration with Gonzalo spatio-temporal point processes (Fig. 1). Tanjona took Ferraz, whom Otso visited in Manaus last year. part to one of the workshops in ZiF with the aim of Jussi developed statistical methods for combining applying these methods in a model of metacommunity individual-level data and population-level data dynamics of wood-decaying fungi. (see also the EBFB project on page 28). Henjo and Otso and Henjo taught in a 5-day PhD Workshop Maria made progress with analysing the movement entitled “Movement Ecology: Analyzing Animal behaviour of elephants and eagle owls, respectively. Movement Data” at the Hebrew University of Jerusalem. Markku finished the development of statistical The field excursion to the desert was a hot event (Fig. 2)!

Collaborators Campioni L., Delgado M., Lourenço R., Bastianelli Prof. Juha Merilä, Department of Biological and G., Fernandez N., Penteriani V. (2013) Individual Environmental Sciences, Helsinki University and spatio-temporal variations in the home range Dr. Vesa Selonen and M.Sc. Andrea Santangeli (University behaviour of a long-lived, territorial species. Oecologia, of Turku), doi: 10.1007/s00442-012-2493-7. Dr. Gonzalo Ferraz, Smithsonian Tropical Research Gurarie E. and Ovaskainen O. (2013). Towards a general Institute, Manaus, Brazil formalization of encounter rates in ecology. Theoretical Dr. Juan Manuel Morales, Ecotono, INIBIOMA— Ecology, doi:10.1007/s12080-012-0170-4. CONICET, Argentina Karhunen M. and Ovaskainen O. (2012). Estimating Dr. Stephen Cornell, Faculty of Biological Sciences, Leeds population-level coancestry coefficients by an University, UK admixture F-model. Genetics 192, 609-617. Prof. Yuri Kondratiev, Dr. Dmitri Finkelshtein and Dr. Saastamoinen M., Brakefield P.M and Ovaskainen, O. Oleksander Kutovyi, University of Bielefeld, Germany (2012). Environmentally induced dispersal-related life history syndrome in the tropical butterfly, Bicyclus Recent Publications anynana. Journal of Evolutionary Biology 25, 2264-2275. Sundell J., Church C. and Ovaskainen O. (2012). Spatio- de Boer W.F., van de Koppel S., de Knegt H. J. and Dekker temporal patterns of habitat use in voles and shrews J. J. A. (2013). Hibernation site requirements of bats in modified by density, season, and predators. Journal of man-made hibernacula in a spatial context. Ecological Animal Ecology 81, 747-755. Applications, doi:10.1890/12-0539.1.

27 European Boreal Forest Biodiversity (EBFB)

Otso Ovaskainen Maria Delgado Project leader Post doc

Jussi Jousimo Juri Kurhinen PhD -student Project coordinator

IT-designer: Evgeniy Meyke Tecnhician: Coong Lo

his project (funded by an Academy of Finland at the Mekrijärvi Biological Station and focused research grant for 2011-2015) aims at putting on collecting and analyzing winter track data (Fig. together a broad-scale and long-term database 1). The second workshop was a PhD course held in Ton European Boreal Forest Biodiversity (EBFB). The St. Petersburg and focused on mathematical and EBFB database shall cover the boreal zones of European statistical modelling of ecological data (Fig. 2). In Russia and Finland and include a description of the addition, Otso and Juri had a memorable trip to the environmental changes (e.g. in terms of forest structure Komi Republic, visiting the Institute of Biology in and climatic conditions) that have occurred over the Syktyvkar and the Pechoro-Ilychskii Nature Reserve. past 50 years as well as the population dynamic and In 2012 we compiled a large fraction of the phenological responses of various taxonomical groups. targeted Finnish data (winter track counts on game The project is based on collaboration between Russian animals; GPS data on wolf, bear, moose, wild forest and Finnish participants. reindeer and lynx; small mammal data; many kinds The motivation for compiling the EBFB database of data on flying squirrel). We acquired pilot versions is that it makes it possible to address many basic and of Russian data, such as coarse-grained data on winter applied scientific research questions of fundamental track counts from European Russia (Fig. 3) and relevance for the fields of ecology, conservation biology phenology data (e.g. arrival dates of birds, flowering and climate change research. In particular, we aim to dates of plants, many kinds of weather data for use this database to disentangle the roles of abiotic and 1960-2012) from the Kivach Nature Reserve. Jussi biotic factors shaping species abundance, distribution focused on modelling the spatio-temporal patterns of and spatio-temporal population dynamics. population abundance whereas Maria took part in the analysis of the phenology data. Highlights of the year

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

28 MRG -ANNUAL REPORT 2012 Mathematical ecology

Fig. 1. Participants of Mekrijärvi meeting on winter track Fig. 2. Eli Gurarie (who used to be a post doc in our data. group) taught computer exercises in the PhD course which we organized for Russian PhD students in St. Petersburg.

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 Recent Publications The EBFB project has 26 Russian and Finnish partner Korpela K., Delgado M., Henttonen H., Korpimäki E., organizations, see http://www.helsinki.fi/science/ Koskela E., Ovaskainen O., Pietiäinen H., Sundell J., metapop/EBFB/participants.html Yoccoz N. G. and Huitu O. (2013). Non-linear effects of In particular, we would like to acknowledge here Prof. climate on boreal rodent dynamics: mild winters do not Harto Lindén, Dr. Ilpo Kojola and Dr. Jyrki Pusenius negate high-amplitude cycles. Global Change Biology, (Finnish Game and Fisheries Research Institute), Dr. DOI: 10.1111/gcb.12099. Ilpo Hanski (Finnish Museum of Natural History), J. Kurhinen, E. Kulebjakina, E. Zadiraka, V.Mamontov, Prof. Andrei Brodsky and Prof. Vladimir Levchenko E. Muravskaya, I. Hanski. (2012). Distribution of the (St. Petersburg State University), Dr. Marina Yakovleva Siberian flying squirrel (Pteromys volans L.) in taiga (Kivach Nature Reserve) prof. Sergei Kochanov and isthmuses between Baltic and White Sea regions. Acta Dr. Eugene Poroshin (Komi Science Centre), Dr. Zoologica Lituanica, 2011, Vol 21(4): 306-310. Andrei Kuprijanov (Pechoro-Ilychskii Nature Reserve) and director Anton Bersenev (Hunting and wildlife Ministry of Russia).

29 Biodiversity conservation informatics

Atte Moilanen Jussi Laitila Project leader Researcher

Enrico Di Minin Tuuli Toivonen Post doc Researcher

Federico Montesino Pouzols Researcher Peter Kullberg Joona Lehtomäki PhD -student PhD -student

Undergraduate students: Ninni Mikkonen, Timo Vaahtoranta

ur group develops concepts, theory, algorithms, methods and software for the needs of conservation prioritization – either Ospatially 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. Fig. 1. The cover of the RobOff v1.0 manual – our new Understanding the implications of uncertainty to software, which is targeted for the allocation of different conservation decision making also is relevant for us. We conservation actions (e.g. habitat management or restoration) work on applications together with collaborators on all and biodiversity offsetting. Cover artwork: Aija Kukkala. continents.

30 MRG -ANNUAL REPORT 2012 Systematic conservation planning

Setting of objectives investigations about knowledge gaps in global conservation prioritization, aiming to utilize the GIS data base in Data Preparation of the a PhD project starting in 2013. Timo preparation ecological model Vaahtoranta has been working on an Preprocesssing of data MSc thesis about the classification and taxonomy of conservation strategies, Monitoring with the intention of soon expanding this work into a PhD thesis. Computational Spatial prioritization GEDA members who started last year analysis have been busy with ongoing research. Dr Jussi Laitila has continued his work on the Priority ranking theory and methodological background Interpretation of spatial ecology and conservation Postprocessing ”Groundtruthing” prioritization. His recent work has ranged from the analysis and comparison Verification of conservation planning paradigms to mathematical tools for spatial decision making. Some of his current research Recommendation for interests include the conceptual action background of conservation biology, the optimality of conservation decision Fig. 2. A schematic presentation of spatial prioritization process, which making and the analysis of alternative the team attempts to improve either on conceptually or methodologically. conservation strategies. Dr. Federico Figure adopted from Lehtomäki and Moilanen (submitted). Montesino Pouzols has concentrated on developing new methods for spatial conservation planning (within the Zonation Highlights of the year framework) and developing the new RobOff framework and software. He visited ACERA and the University of This year has been the second of our ERC project GEDA Melbourne last summer, together with Atte Moilanen, (Global Environmental Decision Analysis). GEDA is and has been collaborating on various projects in funded by a European Research Council (ERC) starting Finland, New Zealand and Australia. grant, and it provides our base-funding for 2011- Locally in Finland, the MetZo-project, “Ecologically 2015. GEDA focuses on the development of concepts, based decision analysis in the implementation of the analyses, software and large-scale applications of spatial South-Central Finland forest biodiversity program”, conservation planning. started in 2010. We develop ecological models for forest This year, GEDA and the renewed CoE have brought and peatland environments together with stakeholders, several new members to our group. Following receipt of including much of the Finnish environmental a PhD from the University of Kent, Dr Enrico Di Minin administration. In this project methods developed in started in April with a focus on conservation, economics, our group (Zonation; RobOff) are fitted and applied and societal impacts of conservation. He has a strong to conservation decision making in Finland. In this background in modeling, economics and GIS, and a context, Joona Lehtomäki continues working on his personal interest in conservation in Africa. Dr Tuuli PhD thesis on application of spatial prioritization Toivonen (senior lecturer in geo-informatics) started methods to planning of forest conservation and in August. She primarily works on understanding data management in Finland, half-time employed by the for spatial conservation and on the combination of Finnish Environment Institute. This year, national land-use zoning and urban planning with biodiversity regional forest analyses were completed and they are analyses. Since the spring, Tuuli has been guiding the being taken onto the field. Also, the national peatland collation of high-resolution global GIS data for the analysis has progressed several rounds and “final” purpose of spatial modeling and conservation planning. results are expected next spring. The next major This work has been assisted by Johanna Kuusterä (MSc objective in this project is the integration of forest and in geographhy), who is replacing the GEDA research peatland conservation planning. secretary Aija Kukkala during her maternity leave. In Also under the MetZO umbrella,Ninni Mikkonen the autumn, Peter Kullberg has started preliminary finished her MSc thesis work “Identification of top

31 priority areas and management landscapes from Our oldest software product, Zonation (since 2004), a national Natura 2000 network”. In this work, a has improved a lot this year. Dr Federico Montesino prioritization was developed across the Finnish Natura Pouzols has implemented great improvements into both 2000 network, with applications for the targeting the core and user interface of the Software. The V3.1 of habitat protection, maintenance and restoration. manual was printed and distributed spring 2012, and Currently, Ninni works at the Natural Heritage Services presently we are working towards version 4.0. During of Metsähallitus (Finnish forest and park service) to 2012, Victoria Veach has put in major effort reviewing help decide expansions of the Finnish forest protection and revising the documentation of Zonation 4 as well area network. as other documents, including www pages, teaching The group leader Atte Moilanen has mainly worked material, etc. with initiation of new projects together with the Next year we expect (i) to continue the development expanded group. Some topics presently of interest for of software Zonation and Roboff, (ii) the completion him include the concept of conservation value, methods of development of the global high-resolution GIS data for habitat management and restoration, development base for global analyses of conservation priority , (iii) of new operational approaches for multi-action initiation of a major new project for the analysis of conservation planning, development of methods and conservation strategies, which was 12/2012 kindly analyses relevant for the Nagoya CBD resolution, and funded by the Kone Foundation (iv), progress for both analysis of alternative conservation strategies. This year, forests and peatlands at the national planning stage, (v) Atte Moilanen together with Jussi Laitila and Federico expansion of the group by two PhD students (Peter Montesino Pouzols taught two courses “Concepts Kullberg, Timo Vaahtoranta and Victoria Veach) to and principles of spatial conservation planning”, formally start during the spring. and “Methods and software of spatial conservation prioritization”, both 3 ECTS.

Collaborators by professor Hugh Possingham. National Prof. Bill Sutherland and Dr Lynn Dicks of Conservation Evidence, University of Cambridge. The South-Central Finland Forest Biodiversity Programme Prof. John Leathwick, Department of Conservation, New (METSO) and its ecological decision analysis umbrella Zealand. project, MetZo, chaired by Dr Mikko Kuusinen at the Lab of Prof Chris Thomas, University of York. Ministry of Environment. The MetZo project involves the Finnish Ministry of Environment, the Ministry of Agriculture and Forestry, the Finnish Environment Recent key publications Institute, the Forest and Park Service (Metsähallitus), the Pouzols F.M., Burgman M.A., and Moilanen A. (2012). Finnish Forest Research Institute, universities, forestry Methods for allocation of habitat management, centers, environmental centers (ELY). maintenance, restoration and offsetting, when The NatNet LIFE+ project for improving the conservation conservation actions have uncertain consequences. area network of South-Western Lapland. (involves e.g. Biological Conservation, 153: 41-50. the Forest Research Institute, Environment center & Laitila J. and A. Moilanen. (2012). Use of many low-level Forestry center Rovaniemi / Lapland) conservation targets reduces high-level conservation Prof. Janne Kotiaho, Prof. Mikko Mönkkönen, Dr Adriano performance. Ecological Modelling 247: 40-47. Mazziotta, and Mr Santtu Kareksela, University of Kujala H., Burgman M.A., and Moilanen A. (2013). Jyväskylä. Treatment of uncertainty in conservation under climate change. Conservation Letters, early doi: 10.1111/j.1755- International 263X.2012.00299.x. ACERA, Australian Centre of Excellence in Risk Analysis, Kukkala A. and A. Moilanen. (2013). The core concepts directed by professor Mark Burgman of spatial prioritization in systematic conservation CEED, Australian Centre of Excellence in Environmental planning. Biological Reviews, doi: 10.1111/brv.12008. Decisions, University of Queensland, Australia, directed

32 MRG -ANNUAL REPORT 2012 Systematic conservation planning

33 Conservation effectiveness

Mar Cabeza Henna Fabritius Project leader Anni Arponen PhD -student Post doc

Johanna Eklund Silvija Budaviciute PhD -student PhD -student Laure Zupan Ricardo Rocha PhD -student PhD -student

Undergraduate student: Helena Uotila

rotected areas cover twelve percent of the Earth’s Highlights of the year terrestrial surface, yet they are insufficient. They are inefficient in covering biodiversity, and they Johanna visited Lauren Coad’s team (Oxford) areP ineffective in protecting it. Population declines to discuss the deforestation project and future as well as illegal deforestation in protected areas collaboration in investigating the links between worldwide indicate that reserves are not functioning. governance and effectiveness. Ricardo and Johanna Part of the problem stems from a history of ad-hoc have been analyzing deforestation in Madagascar and 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; © Helena Uotila Examining the effectiveness of local management practices on the population viability of a butterfly in Fig. 1. Investigating deforestation causes in Madagascar: Finland. Resource mapping activity at a Malagasy village.

34 MRG -ANNUAL REPORT 2012 Systematic conservation planning

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.

Helena spent three months in the field conducting conserving farmland biodiversity in collaboration with interviews, to try to understand the social drivers of the Finnish Environment institute. Henna has collected some of the patterns encountered (Fig. 1.). The visit and analyzed institutional and ecological data to of Mr. Mamy Rakotoarijaoana, the Ranomafana demonstrate the deficiency of short-term management National Park director (Madagascar) initiated an plans for her study species, the endangered butterfly M. interesting collaboration from which we expect new diamina. research angles. Silvija has concentrated on sorting and identifying the massive ant samples she got from Madagascar in late 2011, and has also measured Collaborators morphological characters of over 2000 individuals. These will be used to address the importance of Lauren Coad, ECI, University of Oxford, UK functional traits and trait groups in performing Neil Burgess, WWF USA particular ecosystem functions in disturbed habitas. Mamy Rakotoarijaona, Madagascar National Parks Ricardo has now completed one and a half years of Wilfried Thuiller, CNRS, France fieldwork in Central Amazon, Brazil. His PhD aims Tarmo Virtanen, Dept Env Sciences, University of Helsinki at assessing the long-term impacts of deforestation Aino Juslén, Andrea Santangeli, Finnish Museum of upon tropical biodiversity, using bats as a study Natural History model (Fig. 2.). Laure has started to evaluate a facet Mikko Kuussaari, Saija Sirkiä Finnish Environment of biodiversity that is often ignored in effectiveness Institute, Natural Environment Centre Riikka Paloniemi, Jukka Similä, Finnish Environment evaluations: the representation of evolutionary history Institute, Environment Policy Centre in conservation areas. She found that within Europe, protected areas fail to cover important areas of high phylogenetic diversity. Finnish conservation policies Publications are also receiving more of our attention. What started Arponen A., et al. Improving conservation planning of as a paper discussion in our regular journal club, semi-natural grasslands: integrating connectivity into has turned into a rather large collaborative project. agrienvironment schemes. Biological conservation, Anni is leading this collaboration, which looks into under revision. the impact of conservation investments on trends in Zupan L. Cabeza M., et al. A conservation dilemma: extinction risk. Thus far we have put substantial effort spatial mismatch of phylogenetic diversity across three into collecting comprehensive datasets, which we vertebrates groups and protected areas in Europe. are now beginning to analyse. Anni is also leading Proceedings of the Royal Society B., under revision. a project on the effectiveness of Finnish policies in

35 Climate change

Mar Cabeza Heini Kujala Project leader Laura Meller Phd -student PhD -student

Maria Triviño Raquel Garcia Phd -student Astrid van Teeffelen PhD -student Post doc

Undergraduate student: Antti Takolander

ur team covers a broad range of topics in when using atlas data. She also reviewed how different relation to climate change and biodiversity. types of uncertainties are addressed in climate change Our research interests range from methods to research, and demonstrated that despite uncertainties, Oassess the observed impacts on species and conservation scientifically robust solutions both in modelling of areas, to developing and improving tools to project future impacts and in spatial conservation prioritization future impacts. Furthermore we develop approaches can be achieved. Maria showed that climate remains the to include future projections in conservation planning. most important factor for modelling the distribution We are interested in addressing the plethora of of species in the Iberian Peninsula. She also found that uncertainties at various levels, in particular those related the species most exposed to future climate changes are to predicting impacts and decision-making processes. not those with most sensitive to it, according to their 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 © Ricardo Rocha findings relate largely to uncertainty in climate change research and have resulted in three papers. She Fig. 1. Whinchat, Saxicola rubetra, one of the species investigated how spatiotemporal changes in survey projected to be most exposed and most sensitive to global. effort affect measurements of observed range shifts change in the Iberian Peninsula (Triviño 2012).

36 MRG -ANNUAL REPORT 2012 Systematic conservation planning

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.

Collaborators Miguel B. Araújo (National Museum of Natural Sciences, Spain) Neil Burgess, WWF US Mark. A. Burgman, University of Melbourne Jon E. Brommer, University of Turku Sebastiaan Deetman, Netherlands Environment Assessment Agency Wendy Foden, IUCN, Cambridge, UK Thomas Hickler, Goethe University, Germany Andries Hof, Netherlands Environment Assessment Agency Aleksi Lehikoinen, Finnish Museum of Natural History Fig. 2. Maps illustrating the major sources of uncertainties Carsten Rahbek, University of Copenhagen , Denmark in projections of species turnover for the year 2050. Wilfried Thuiller, University of Grenoble, France Variation is larger among different Bioclimatic Envelope Models (BEM) than emissions scenarios (SRES), although Key publications some locations in Africa also show disagreement between climate models (GCMs) (see Garcia et al. 2012) 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 traits and threat status, with few exceptions (Figure vertebrates. Global Change Biology 18:1253–1269 1). Raquel’s work has focused on climate change Kujala, H. (2012). Climate change, species range shifts and impacts on African vertebrates She has investigated the uncertainty – a new era of conservation planning. PhD methodological uncertainties in results from species thesis, Faculty of Biological and Environmental Sciences, distribution models (SDM). She found most of the University of Helsinki. Unigrafia Oy, Helsinki. variation in projections to result from the choice of Kujala, H., Burgman, M.A. and Moilanen, A. Treatment modelling algorithm, and compared different ways to of uncertainty in conservation under climate change. generate consensus around the projections (Figure (2013). Conservation Letters, doi: 10.1111/j.1755- 2). She is currently looking at whether species at risk 1263X.2012.00299.x identified by SDMS match with intrinsic traits that Kujala, H., Moilanen, A., Araújo, M.B. and Cabeza, M. (2012). Conservation planning with uncertain climate potentially influence the species’ response. change projections. PLOS ONE (in press) Much of the work produced sits within the EU-FP7 Kujala, H., et al. (2013) Range margin shifts of birds project RESPONSES (2010-2013), for which Mar leads revisited — the role of spatiotemporally varying survey the Biodiversity work package. RESPONSES investigates effort.Global Change Biology, 19:420-430. EU policy action on climate change covering a range Meller, L., Barbet-Massin, M., Deetman, S., Hof, A., Pironon, of sectors and communicating findings to the EU. In S., Thuiller, W., & Cabeza, M. (2012). Joint impacts of August, Mar gave a well-received keynote speech at the climate change and bioenergy production on the state Nordic Climate Adaptation conference, summarizing and conservation of European birds. RESPONSES project the outcomes of RESPONSES project. We also produced deliverable 5.3a (p. 30 pp.). a policy brief that stressed the gaps between science and Triviño, M., 2012. Global change impacts and conservation policy and also highlighted the expected reduction in priorities in the Iberian Peninsula. PhD dissertation European conservation effectiveness due to climate in Ecology. University Rey Juan Carlos and National Museum of Natural Sciences, Madrid. change. We organized a workshop with biodiversity Van Teeffelen, A., Lung, T., Meller, L., Vermaat, J., Lavalle, experts in Ehrenberg, Germany to assess cross- C., & Cabeza, M. (2012). EU Biodiversity policy in times sectoral opportunities and threats from mitigation of climate change. RESPONSES policy update No. 1. and adaptation. The outcomes are being combined Van Teeffelen, A.J.A., Vos, C.C. & Opdam, P. (2012). with a literature review and a model analysis. Laura is Species in a dynamic world: Consequences of habitat preparing a manuscript exploring the relative impacts network dynamics and conservation planning. Biological of climate change and bioenergy, thereby addressing the Conservation, 153, 239-253.

37 Supporting personnel

do the laboratory work in the project of wood- I 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-Kovero Laboratory technician

y main task is to take care of the MGlanville fritillary (Melitaea cinxia) laboratory populations and assist researchers in their field and laboratory experiments. I’m working year around at Lammi Biological Station. Suvi Ikonen Coordinator

am taking care of data recording during I 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 Meyke IT designer facilitate the laboratory work of I 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 Nyman Laboratory technician

y tasks in the Glanville fritillary project Minclude 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 Ruokolainen Laboratory technician

38 MRG -ANNUAL REPORT 2012 Supporting personnel

y main duties consist of group’s financial and personnel Madministration as well as international project management. I am also taking care of various other research supporting tasks.

Viia Forsblom Research secretary am organizing and supervising large I 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 Sami Ojanen group. Project coordinator

am a lab technician in Mikko Frilander’s group. My main I 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.

Marja-Leena Peltonen Laboratory technician I work in Anna-Liisa Laines group of I 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 Raveala Research secretary

do laboratory work in the Glanville fritillary project. For Inow 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. Suvi Saarnio Laboratory technician

y main tasks in the Biodiversity Con- Mservation 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 Aija Kukkala documentation of software. On maternity leave. Research secretary

39 do laboratory work in the Glanville fritillary project. I am in Icharge 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. Pia Välitalo Technical assistant

have been editing the Zonation and Roboff Imanuals for Atte. I have also been helping with data pre-processing for Mar’s group.

Victoria Veach Technical assistant

am taking care of personnel and financial Iadministration tasks and working with GIS database building and other tasks.

Johanna Kuusterä Research secretary

40 MRG -ANNUAL REPORT 2012 Synopsis of the year 2012

Publications Theses Articles and book chapters

Honours, awards and memberships

Visitors

Teaching

Budget

Annual meeting in St Petersburg

PhD theses

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

ll species are adapted to certain climatic conditions, dynamics can lead to misguided and potentially Aoutside of which they cannot survive. Changes inefficient conservation decisions. The results presented in the climatic environment therefore force species here show that modelling future impacts using so- to either adapt to the new conditions or move to areas called niche modelling techniques, despite their inherent where suitable conditions are still present in order uncertainties, can provide useful information about to avoid extinction. Several studies have shown that how species distributions and conservation statuses species from various taxa are currently moving their will be affected by climate change. For example ranges polewards and to higher elevations to keep previously created models for Finnish breeding birds up with shifting climate regimes. However, species predicted well recently observed changes in species differ widely in their dispersal abilities. In addition, distribution sizes. More importantly, the observed natural landscapes are becoming increasingly changes seem to match best with predictions that human-dominated, further hindering dispersal by follow the most extremeclimate change scenario. A decreasing permeability. anthropogenic climate key factor for successfully measuring and predicting change is therefore expected to become one of the climate change impacts are good monitoring data, the major drivers of species extinctions by the end of the role of which should be more widely acknowledged by 21st century. decision-makers. uncertainty in climate change research Species range shifts are problematic in conservation is pervasive and cannot ever be entirely eliminated. planning, because dynamic biodiversity patterns This work offers tools to assist in both spatial hamper our ability to identify priority areas for prioritization and decision making when scarce protection. because protected area networks are conservation resources need to be allocated under geographically fixed, climate change may also drive uncertain future conditions. The findings of this thesis species out of reserves, foiling past conservation strongly encourage using proactive approaches that efforts. in this thesis the different risks and account for future impacts. The results also suggests opportunities of conducting conservation planning that while striving to reduce epistemic uncertainty under climate change are investigated. This research is important in climate change and conservation focuses on the uncertainties that arise from working research, other sources of uncertainty such as socio- with unknown future events, technical challenges political factors or volitional human behaviour might of observing and predicting species range shifts, and constitute far larger determinants of successful using (or ignoring) information about future impacts conservation actions, and therefore merit stronger in conservation planning. focus in research. The major findings of this thesis are that climate change is already rapidly reshaping species Supervisor: Mar Cabeza distributions in Finland and that ignoring future

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

any animal species live in societies in which individual fitness. For example, in territorial species Mnearby conspecifics are vital elements of their where at least two social groups –breeders and nonsocial environment, with the nature and quality of their territorial floaters – are recognized, territorial ownership behavioral interactions determining the type of social can lead to holders behaving differently compared to the organization. As a group, birds show a wide range of floating counterpart of the population. For this reason, social organizations where, in some cases, social status social structure is often considered a key determinant of gives priority of access to resources, ultimately affecting population biology, influencing fitness, gene flow, and

42 MRG -ANNUAL REPORT 2012 Synopsis of the year

spatial pattern and scale. Nonetheless, nonsocial factors the home range size (mean HR size ~ 220 ha) appears (e.g., environmental condition, food supply) can also to be a direct consequence of landscape structure rather affect behavioral interactions, individual relationships, than prey abundance available across the study area. and, ultimately, social organization. In the present Across the mosaic of territories settled by owls, females thesis, we studied the behavioral differences between - the sex that experiences less social constriction - are individuals of different social status; we focused in those which exhibit wider home ranges that overlap particular on the analysis of habitat selection, space to a greater extent with those of their neighbors. use behavior and movement patterns of breeders and Nevertheless, within the boundaries of their home nonterritorial eagle owls (Bubo bubo). ranges, adults’ behavioral decisions were significantly The focal radio-tracking of breeders and affected by nonsocial factors such as the biological nonterriotrial floaters during 8 years demonstrated needs and individual identity. Similarly, external cues that owls perform different behavioral strategies in like the lunar cycle (Chapter 3), act to regulate the time relation to different life cycle stages, social status and and effort that owls allocated to social (communication) the behavioral trait under study. These observations or physiological (feeding) activities. emphasize the existence of more structured inter- A key finding demonstrated here is that individual relationships than expected. Moreover, nonterritorial floaters show a tremendous capacity previous investigations of social interactions (vocal to adapt their behaviour to their immediate needs and visual communication) support the importance and social and physical surroundings. As with other of territoriality and social dominance on owls’ territorial species, floating owls show cryptic behaviour, behavioral decisions. Our results indicate a scenario living in a parallel “underworld” where individuals in which both social and nonsocial factors seem to make decisions while considering social constraints affect the behavioral mechanisms that regulate habitat (Chapter 1-2), acquired experience (Chapter 4) and selection, space use and movement behavior in different landscape features. At the end of their natal dispersal, ecological contexts. In contrast to our initial predictions, the most likely fate for our floaters was to settle close trophic resource abundance in our study area does to the natal population while awaiting circumstances not correlate directly with owls’ space use behavior. that would offer greater reproductive opportunities. However, the large abundance of the staple prey across In conclusion, the study of the relationships between this area, due primarily to management and release of animals and their environment is a field where ecology rabbits (the study area serving as game reserve), might and behavior are tightly intertwined. In my opinion, and actually favor a high density of conspecifics over a as stressed in the present study, social organization is a reduced area (40 breeding pairs/100 km2) by relaxing key determinant of population biology with important environmental constrictions like resource competition implications on spatial processes. (e.g. food). In line with this prediction, we show that territory holders (Chapter 5) occupy reduced home Co-Supervisor: Maria Delgado ranges of high quality for reproduction. Surprisingly,

MSc theses

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

n my MSc thesis I studied the effects of habitat The experiment was carried out in 2010-2012 and Ifragmentation on fitness and genetic diversity in it was part of a more extensive study which aims to find a wind-pollinated plant species Plantago lanceolata. out how well different proxies account for plant’s life- Human induced habitat fragmentation is one of the most time reproduction success. 40 populations representing severe problems for survival of many species. Habitat the natural variation of the system in Åland Islands fragmentation reduces population sizes and isolates were chosen for the study. The study speciesPlantago populations from each other. In smaller and isolated lanceolata is a diploid perennial. It is self-sterile and populations the risk of inbreeding depression increases wind-pollinated plant that also proliferates clonally. It and that affects fitness of the populations. The effects has a tendency of forming a soil seed bank where the of habitat fragmentation on plant fitness and genetic seeds may survive viable for a number of years. The variation may also depend on plant characteristics such population in Åland Islands is naturally fragmented and as life span, mating system and rarity. located at the Northern edge of this species range .

43 I studied if the degree of isolation and size of the populations in Åland Islands don’t seem to be seriously population have any effect on seed weight, germination affected by the fragmented population structure and plant size. The only significant correlation was and the reason for that may be found from the plant between population connectivity and plant size, plants characteristics such as clonal reproduction and the in isolated populations were smaller than in populations formation of extensive soil seed bank. that were better connected to other populations. The state of Plantago lanceolata in Åland seem to AMOVA analysis of the genetic data revealed be fairly good at the moment, but it is advisable to keep that the amount of inbreeding in these populations is track on the state of the population in future because fairly high. Still the populations did seem to have some the negative impacts of inbreeding may arise if the gene flow among them, because the populations didn’t population suffers from stressful conditions. seem to be strongly differentiated from each other. The inbreeding didn’t seem to affect viability of the seeds Supervisors: Anna-Liisa Laine (germination percentage being roughly at the same Saskya van Nouhuys level as in continuous populations. Plantago lanceolata

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

abitat fragmentation is one of the major threats to Phenotypes of both species showed some sensitivity Hspecies diversity around the world. It increases the to the size and isolation levels of local populations, distance between remnant habitat patches and decreases but overall the detected effects were relatively small. their size. As a result, the population sizes of the species Individuals from small local population of P. lanceolata inhabiting the fragments is reduced and their isolation were on average smaller than individual from large is increased. Isolation and small population size can lead populations (F(1, 43.2) = 5.04, p = 0.031). Leaf size of to inbreeding and loss of genetic variation. Inbreeding V. spicata was smaller in highly connected populations depression is expected to cause reductions in the fitness (F(1, 33.6) = 9.27 p = 0.005). The effects of populations traits of the species inhabiting the remnant fragments. size to leaf size of V. spicata was different within stress Habitat fragmentation alters also the environmental groups (Interaction: F(1,117) = 4.04, p = 0.029). In conditions and species interactions of the populations stressed group leaf size was largest in small populations, living in remnant patches. This can lead to changes in but in non-stressed leaf size was largest in large the selection pressure of the fitness traits and further to populations. adaptive evolution. None of the detected effects could be unambiguously In this thesis, the effects of habitat fragmentation accounted to be caused by inbreeding depression. on two plant species, wind pollinated P. lanceolata and The effects were more likely caused by environmental insect pollinated V. spicata, are studied with a common factors that are correlated with population isolation and garden experiment. Both species are perennial herbs size. Larger size of V. spicata in isolated populations living in fragmented environment in South-West could also be caused by adaptation to pollen limitation Finland. From both species, four individuals from 40 in isolated populations. This study has shown that different populations with different size and isolation fragmentation does not necessarily lead to inbreeding levels were used. Two treatments stress and reduced depression in small and isolated populations and species pollination were applied. In the end of growing season can be adapted to living in fragmented conditions. fitness related traits from the plants were measured. Interpopulation gene flow, purgin, and certain species Principal components were extracted based on the characters can be responsible for the low levels of measurements. Components of P. lanceolata were: inbreeding depression. plant size, number of flowers and leaves, and see set. Components of V. spicata were: leaf size and number Supervisors: Anna-Liisa Laine of leaves. Effects of isolation and population size to the Saskya van Nouhuys principal components were analysed with linear mixed models.

44 MRG -ANNUAL REPORT 2012 Synopsis of the year

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

The purpose of this master’s thesis is to study nature conservation prioritization. Data consisted of 68 values within the Finnish national Natura 2000 network Natura 2000 habitat types and their state of naturalness on state owned land. The six goals of this work were and representativeness. Zonation took into account the achieved: 1) Areas with most nature values were rarity, quality, importance, threat status, biodiversity identified by prioritizing habitats of Natura 2000 directive value, congruity and connectivity of these habitat types. (92/43/EU) within. Areas with high nature value were As a result software produces a map of conservation usually in very natural state and had good connectivity priorities and associated quantitative information, to other similar places, or they were spots of some very which facilitate identification of areas with most rare nature types. 2) It was found out that data used was considerable nature values. These were identified both suitable for identifying conservation values, 3) find out ocularly and with Zonation software. the suitability of Zonation software in conservation area Analyses were done at two levels: all habitat types management and maintenance planning and 4) find together and in subgroups following division to major out how results will change if conservation status is habitat types, such as coastal environments, inland taken into account. As an addition to these 5) the most waters, meadows, alpine habitats, peat lands, rocky considerable areas with high conservation value were areas and forests. Results showed that connectivity identified and 6) “Zonation software in a nutshell” was increased aggregation of areas with high nature values produced in Finnish to assist Finnish state officials to use and weighting spread them. Hierarchical analysis was the software for conservation purposes. These results used to find out how nature values changed when will help Metsähallitus (The Finnish Forest and Park the conservation status of the areas were taken into Service) - Natural Heritage Service - to target resourcing account. The results of hierarchical analysis show that of habitat management and restoration in and around conservation status changed the results a lot. Difference the areas with most considerable nature values. It is between main analysis and hierarchical analysis was essential to sustain these areas and their values so that much greater than when taken into account connectivity their nationwide importance can be maintained into the of feature weights. Hierarchical comparison revealed future. that many areas with considerable high nature values Data used in this study covered areas that were areas are not presently strictly protected. classified as Natura 2000 habitats according to European Union Council Directive 92/43/EEC. Analyses were Supervisor: Atte Moilanen done by using Zonation software, a tool for spatial

Articles and book chapters planning for biodiversity and development in the urban fringed. Landsc. Urban Plann. 108:140-149. Bonte D. and Saastamoinen M. (2012). Dispersal Alagador D. A., Triviño M., Cerdeira O., Bras R., Cabeza syndromes in arthropods: butterflies and spiders as a M., and Araújo M. B. (2012). Linking like with like: model system. in M. Baguette, T. Benton, J. Bullock, and optimising connectivity between environmentally- J. Clobert, editors. Dispersal and Spatial Evolutionary similar habitats. Landsc. Ecol. 27:291-301. Ecology. Oxford University Press, Oxford. Arponen A. (2012). Prioritizing species for conservation Bonte D., Van Dyck H., Bullock J. M., Coulon A., Delgado planning. Biodivers. Conserv. 21:875-893. M. d. M., Gibbs M., Lehouck V., Matthysen E., Mustin K., Arponen A., Heikkinen R. K., Paloniemi R., Pöyry J., Similä Saastamoinen M. A. K., Schtickzelle N., Stevens V. M., J., and Kuussaari M. (in press). Improving conservation Vandewoestijne S., Baguette M., Barton K., Benton T. G., planning for semi-natural grasslands: integrating Chaput-Bardy A., Clobert J., Dytham C., Hovestadt T., connectivity into agri-environment schemes. Biol. Meier C. M., Palmer S. C. F., Turlure C., and Travis J. M. Conserv. J. (2012). Costs of dispersal. Biological Reviews. 87:290- Arponen A., Lehtomäki J., Leppänen J., Tomppo E., and 312. Moilanen A. (2012). Effects of connectivity and spatial Calandra R. R. T., Deisenroth M. P. & Pouzols F. M. 2012. resolution of analyses on conservation prioritization Learning Deep Belief Networks from Non-stationary across large extents. Conserv. Biol. 26:294-304. Streams. Pages 379-386 in Artificial Neural Networks Bekessy S. A., White M., Gordon A., Moilanen A., and Machine Learning – ICANN 2012. Springer Berlin McCarthy M. A., and Wintle B. A. (2012). Transparent Heidelberg.

45 Campioni L., Lourenco R., Delgado M. d. M., and Penteriani Hanski I. (2012). Eco-evolutionary dynamics in a changing V. (2012). Breeders and floaters use different habitat world. Year in Ecology and Conservation Biology. 1249:1- cover: should habitat use be a social status-dependent 17. strategy? Journal of Ornithology. 153:1215-1223. Hanski I. (2012). Eco-evolutionary dynamics in a changing Campioni L. D. M., Lourenço R., Bastianelli G., Fernandez world. Pages 1-17 in R. S. Ostfeld and W. H. Schlesinger, N., Penteriani V. (2013). Individual and spatio-temporal editors. The Year in Ecology and Conservation Biology. variations in the home range behaviour of a long-lived, Annals of the New York Academy of Sciences. territorial species. Oecologia. doi: 10.1007/s00442- Hanski I. (2012). Metapopulations and spatial population 00012-02493-00447. processes. Oxford Bibliographies in Ecology. http://www. de Boer W. F., van de Koppel S., de Knegt H. J., and Dekker oxfordbibliographies.com/. J. J. A. (2013). Hibernation site requirements of bats in Hanski I. (2012). Spatial structure and dynamics in the man-made hibernacula in a spatial context. Ecol. Appl.: Glanville fritillary (Melitaea cinxia) metapopulation. doi:10.1890/1812-0539.1891. Pages xxviiii-xxxi in J. Clobert, Baguette, M., Benton, De Jong M., Collins S., Beldade P., Brakefield P., and Zwaan T.G. and Bullock, J.M. , editor. Dispersal Ecology and B. (2013). Footprints of selection in wild populations Evolution. Oxford University Press, Oxford. of Bicyclus anynana along a latitudinal cline. Mol. Ecol. Hanski I. (2012). A tribute to the ERC-long live basic 22:341-353. research. EMBO Rep. 13:474-474. Dees M. W., Somervuo P. J., Lysøe E., Aittamaa M., Hanski I. and Kuuluvainen T. (2012). Metsien and Valkonen J. (2012). Species identification and monimuotoisuuden väheneminen jatkuu. Helsingin microarray-based comparative genome analysis of Sanomat. Streptomyces species isolated from potato scab lesions in Hanski I., von Hertzen L., Fyhrquist N., Koskinen K., Norway. Mol. Plant Pathol. 13:174-186. Torppa K., Laatikainen T., Karisola P., Auvinen P., Paulin Di Minin E., Fraser I., Slotow R., and MacMillan D. C. L., Makela M. J., Vartiainen E., Kosunen T. U., Alenius (2013). Understanding heterogeneous preference H., and Haahtela T. (2012). Environmental biodiversity, of tourists for big game species: implications for human microbiota, and allergy are interrelated. Proc. conservation and management. Anim. Cons.: DOI: Natl. Acad. Sci. U. S. A. 109:8334-8339. 10.1111/j.1469-1795.2012.00595.x. Hornett E. A. and Wheat C. W. (2012). Quantitative Di Minin E., MacMillan D. C., Goodman P. S., Escott RNA-Seq analysis in non-model species: assessing B., Slotow R., and Moilanen A. (2013). Conservation transcriptome assemblies as a scaffold and the utility of businesses and conservation planning in a biodiversity evolutionary divergent genomic reference species. BMC hotspot. Conserv. Biol. Genomics. 13:361. Di Minin E. and Moilanen A. (2012). Empirical evidence Karhunen M. and Ovaskainen O. (2012). Estimating for reduced protection levels across biodiversity features Population-Level Coancestry Coefficients by an from target-based conservation planning. Biol. Conserv. Admixture F Model. Genetics. 192:609-617. 153:187-191. Karinen S. H., Saarinen S., Lehtonen R. J., Rastas P., Duplouy A., Iturbe-Ormaetxe I., Beatson S. A., Szubert Vahteristo P. M., Aaltonen L. A., and Hautaniemi S. J. M., Brownlie J. C., McMeniman C. J., McGraw E. (2012). Rule-based induction method for haplotype A., Hurst G. D., Charlat S., O’Neill S. L., and Woolfit comparison and identification of candidate disease loci. M. (2013). Draft genome sequence of the male-killing Genome Med. 4:21. Wolbachia strain wBol1 reveals recent horizontal gene Kool J. T., Moilanen A., and Treml E. A. (in press). transfers from diverse sources. BMC Genomics. 14:20. Population connectivity: recent advances and new Garcia R. A., Burgess N. D., Cabeza M., Rahbek C., and perspectives. Landsc. Ecol.: doi: 10.1007/s10980-10012- Araujo M. B. (2012). Exploring consensus in 21st century 19819-z. projections of climatically suitable areas for African Korpela K. D. M., Henttonen H., Korpimäki E., Koskela vertebrates. Global Change Biology. 18:1253-1269. E., Ovaskainen O., Pietiäinen H., Sundell J., Yoccoz N. Gurarie E. and Ovaskainen O. (2013). Towards a general G. and Huitu O. (in press). Non-linear effects of climate formalization of encounter rates in ecology. Theoretical on boreal rodent dynamics: mild winters do not negate Ecology. doi:10.1007/s12080-12012-10170-12084. high-amplitude cycles. Global Change Biology. Halme P., Vartija N., Salmela J., Penttinen J., and Norros Kos M., Hoetmer A. J., Pretorius Y., Boer W. F., de Knegt H. V. (2012). High within- and between-log variation in the J., Grant C. C., Kohi E., Page B., Peel M., Slotow R., Waal nematoceran community and its physical environment C., Wieren S. E., Prins H. H. T., and Langevelde F. (2012). in decaying aspen trunks. Insect Conservation and Seasonal diet changes in elephant and impala in mopane Diversity. woodland. Eur. J. Wildl. Res. 58:279-287. Hanski I. (2012). Dispersal and eco-evolutionary dynamics Koskinen J. P. and Holm L. (2012). SANS: high-throughput in the Glanville fritillary butterfly. Pages 290-303 in retrieval of protein sequences allowing 50% mismatches. J. Clobert, Baguette, M., Benton, T.G. and Bullock, Bioinformatics. 28:i438-i443. J.M., editor. Dispersal Ecology and Evolution. Oxford Koskinen P., Laine P., Niemi O. A., Nykyri J., Harjunpää University Press, Oxford. H., Auvinen P., Paulin L., Pirhonen M., Palva E. T., and

46 MRG -ANNUAL REPORT 2012 Synopsis of the year

Holm L. (2012). Genome Sequence of Pectobacterium policies. Biol. Conserv. 155:141-148. sp. Strain SCC3193. J. Bacteriol. 194:6004. Martinez J., Duplouy A. M. R., Woolfit M., Vavre F., O’Neill Kosonen L., Schigel D., Setälä R., and Smirnoff I. (2012). S. L., and Varaldi J. (2012). Influence of the Virus LbFV Sieni pusertaa puusta lumihiuksia. Suomen luonto. and of Wolbachia in a Host-Parasitoid Interaction. PLoS 71:10-10. ONE. 7(4). Kraft T. S. and van Nouhuys S. (in press). The effect of host Mattila A., Duplouy A. M. R., Kirjokangas M., Lehtonen R. density and species on superparasitism and sex ratio in a J., Rastas P., and Hanski I. (2012). High genetic load in an gregarious parasitoid. Ecol. Entomol. old isolated butterfly population. Proc. Natl. Acad. Sci. U. Kujala H. (2012). Climate change, species range shifts S. A. 109:E2496-2505. and uncertainty - A new era of conservation planning. Meller L., Barbet-Massin M., Deetman S., Hof A., Pironon Univerity of Helsinki, Unigrafia, Helsinki. S., Thuiller W., and Cabeza M. (2012). Joint impacts of Kujala H., Burgman M. A., and Moilanen A. (2013). climate change and bioenergy production on the state Treatment of uncertainty in conservation under climate and conservation of European birds. RESPONSES Project change. Conservation Letters. doi: 10.1111/j.1755- deliverable: Integrated Activity Report I. part of D 5.3. 1263X.2012.00299.x. Mikkonen N. and Moilanen A. (in press). Identification of Kujala H., Moilanen A., Araújo M. B., and Cabeza M. (in top priority areas and management landscapes from a press). Conservation planning with uncertain climate national Natura 2000 network. Environmental Science & change projections. PLoS ONE. Policy. 27:11-20. Kujala H., Vepsäläinen, V., Zuckerberg, B., and Brommer, Miraldo A., Faria C., Hewitt G. M., Paulo O. S., and J.E. (2013). Range margin shifts of birds revisited – the Emerson B. C. (2012). Genetic analysis of a contact zone role of spatiotemporally varying survey effort. Global between two lineages of the ocellated lizard (Lacerta Change Biology. 19:420-430. lepida Daudin 1802) in south-eastern Iberia reveal a Kukkala A. and Moilanen A. (in press). The core concepts steep and narrow hybrid zone. J. Zool. Syst. Evol. Res. of spatial prioritization in systematic conservation Miraldo A., Hewitt G. M., Dear P. H., Paulo O. S., and planning. Biological Reviews. doi: 10.1111/brv.12008. Emerson B. C. (2012). Numts help to reconstruct the Kurhinen J., Kulebyakina E., Zadiraka E., Mamontov V., demographic history of the ocellated lizard (Lacerta Muravskaya E., and Hanski I. K. (2012). Distribution lepida) in a secondary contact zone. Mol. Ecol. 21:1005- of siberian flying squirrel (Pteromys volans l.) in taiga 1018. isthmuses between Baltic and White sea regions. Acta Moilanen A. (2012). Reserve selection and conservation Zool. Litu. 21:306-310. prioritization. Pages 617-624 in L. Gross, editor. Encyclo- Kvist J., Wheat C.W., Kallioniemi E., Saastamoinen M., I. pedia of theoretical ecology. University of California Press. H., and M. F. (2013). Temperature treatments during Moilanen A. (2012). Spatial Conservation Prioritization in larval development reveal extensive heritable and plastic Data-Poor Areas of the World. Natureza & Conservacao. variation in gene expression and life history traits. Mol. 10:12-19. Ecol. 22:602-619. Moilanen A. (2013). Planning impact avoidance and Laitila J. and Moilanen A. (2012). Use of many low-level biodiversity offsetting using software for spatial conservation targets reduces high-level conservation conservation prioritization. Wildl. Res.: http://dx.doi. performance. Ecol. Model. 247:40-47. org/10.1071/WR12083. Laitila J., Nieminen P. J., Saksman E., and Tylli H.-O. Moilanen A., Anderson B. J., Arponen A., Montesino (2013). Compact and weakly compact composition Pouzols F., and Thomas C. D. (2013). Edge artefacts operators on BMOA. Complex Analysis and Operator and lost performance in national versus continental Theory. 7(1):163-181. conservation priority areas. Divers. Distrib. 19:171-183. Laurentz M., Reudler J. H., Mappes J., Friman V., Ikonen Moilanen A., Leathwick J. R., and West D. W. (2012). S., and Lindstedt C. (2012). Diet Quality Can Play a Development of a systematic, information-based Critical Role in Defence Efficacy Against Parasitoids and approach to the identification of high value sites for Pathogens in the Glanville Fritillary (Melitaea Cinxia). J. freshwater conservation in New Zealand. in P. J. Boon and Chem. Ecol. 38:116-125. P. J. Raven, editors. River Conservation and Management. Leathwick J. R., West D. W., and Moilanen A. (2012). Wiley-Blackwell. Development of a systematic, information-based Moilanen A., Meller L., Leppänen J., Montesino Pouzols approach to the identification of high value sites for F., Kujala H., and Arponen A. (2012). Zonation spatial freshwater conservation in New Zealand. Pages 183-192 conservation planning framework and software V3.1, in P. J. B. a. P. J. Raven, editor. River Conservation and User manual. Management. Wiley-Blackwell Mwakiwa E., de Boer W. F., Hearne J. W., Slotow R., van Liu X., Suzuki A., and Schigel D. (2012). Editorial: The Langevelde F., Peel M., Grant C. C., Pretorius Y., Stigter impact of fungi on other organisms. Mycology. 3:1. J. D., Skidmore A. K., Heitkönig I. M. A., de Knegt Maron M., Hobbs R. J., Moilanen A., Matthews J. W., H. J., Kohi E. M., Knox N., and Prins H. H. T. (2013). Christie K., Gardner T. A., Keith D. A., Lindenmayer Optimization of wildlife management in a large game D. B., and McAlpine C. A. (2012). Faustian bargains? reserve through waterpoints manipulation: A bio- Restoration realities in the context of biodiversity offset economic analysis. J. Environ. Manag. 114:352-361.

47 Nemri A., Barrett L. G., Laine A. L., Burdon J. J., and Thrall 12:1440-1450. P. H. (2012). Population Processes at Multiple Spatial Pouzols F. M., Burgman M. A., and Moilanen A. (2012). Scales Maintain Diversity and Adaptation in the Linum Methods for allocation of habitat management, marginale - Melampsora lini Association. Plos One. 7. maintenance, restoration and offsetting, when Nieberding C. M., Fischer K., Saastamoinen M., Allen C. E., conservation actions have uncertain consequences. Biol. Wallin E. A., Hedenstrom E., and Brakefield P. M. (2012). Conserv. 153:41-50. Cracking the olfactory code of a butterfly: the scent of Pouzols F. M. and Lendasse A. (2012). Adaptive kernel ageing. Ecol. Lett. 15:415-424. smoothing regression for spatio-temporal environmental Niitepõld K. and Hanski I. (2013). A long life in the fast datasets. Neurocomputing. 90:59-65. lane: positive association between peak metabolic rate Pretorius Y., Stigter J. D., de Boer W. F., van Wieren S. E., and lifespan in a butterfly. J. Exp. Biol.: doi:10.1242/ de Jong C. B., de Knegt H. J., Grant C. C., Heitkönig I., jeb.080739. Knox N., Kohi E., Mwakiwa E., Peel M. J. S., Skidmore Nilsson R. H., Tedersoo L., Abarenkov K., Ryberg M., A. K., Slotow R., van der Waal C., van Langevelde F., and Kristiansson E., Hartmann M., Schoch C. L., Nylander Prins H. H. T. (2012). Diet selection of African elephant J. A. A., Bergsten J., Porter T. M., Jumpponen A., over time shows changing optimization currency. Oikos. Vasihampayan P., Ovaskainen O., Hallenberg N., 121:2110-2120. Bengtsson J., Eriksson M., Larsson K. H., Larsson E., and Radivojac P., Koskinen P., Nokso-Koivisto J., Holm L. et Kõljalg U. (2012). Five simple guidelines for establishing al. (in press). A large scale evaluation of computational basic authenticity and reliability of newly generated protein function prediction. Nat. Meth. fungal ITS sequences. MycoKeys. 4:37-63. Saarinen S., Vahteristo P., Lehtonen R., Aittomaki K., Norros V., Penttilä R., Suominen M., and Ovaskainen O. Launonen V., Kiviluoto T., and Aaltonen L. A. (2012). (2012). Dispersal may limit the occurrence of specialist Analysis of a Finnish family confirms RHBDF2 wood decay fungi already at small spatial scales. Oikos. mutations as the underlying factor in tylosis with 121:961-974. esophageal cancer. Familial Cancer. 11:525-528. Nykyri J., Niemi O. A., Koskinen P., Nokso-Koivisto J. Saastamoinen M., Hirai, N., van Nouhuys, S. (2013). Direct P., Pasanen M., Broberg E. M., Pljusnin I., Törönen P., and trans-generational responses to food deprivation Holm L., Pirhonen M., and Palva E. T. (2012). Revised during development in the Glanville fritillary butterfly. Phylogeny and Novel Horizontally Acquired Virulence Oecologia. 171:93-104. Determinants of the Model Soft Rot Phytopathogen Saastamoinen M., Brakefield P. M., and Ovaskainen O. Pectobacterium wasabiae SCC3193. PLoS Pathogens. (2012). Environmentally induced dispersal-related 8:e1003013. life-history syndrome in the tropical butterfly, Bicyclus Ohdachi S. D., Yoshizawa K., Hanski I., Kawai K., anynana. J. Evol. Biol. 25:2264-2275. Dokuchaev N. E., Sheftel B. I., Abramov A. V., Saastamoinen M., Ikonen S., Wong S. C., Lehtonen R., Moroldoev I., and Kawahara A. (2012). Intraspecific and Hanski I. (in press). Plastic larval development in a phylogeny and nucleotide diversity of the least shrews, butterfly has complex environmental and genetic causes the Sorex minutissimus-S. yukonicus complex, based on and consequences for population dynamics. J. Anim. nucleotide sequences of the mitochondrial cytochrome b Ecol. gene and the control region. Mamm. Study. 37:281-297. Schigel D. (2012). Fungivory and host associations of Ovaskainen O. (2012). Pelastetaan vanhat aineistot! Coleoptera: a bibliography and review of research Luonnon Tutkija. 3:63. approaches. Mycology: An International Journal on Ovaskainen O. (2012). Strategies for Improving Fungal Biology. 3:258-272. Biodiversity Conservation in the Netherlands: Enlarging Schigel D. (2012). The impact of fungi on other organisms. Conservation Areas vs. Constructing Ecological Mycology: An International Journal on Fungal Biology. Corridors: An expert report ordered by the Dutch Council 3:1-1. for the Environment and Infrastructure. An expert report Schigel D. S. (2012). Fungivory of saproxylic Coleoptera: ordered by the Dutch Council for the Environment and the mystery of rejected polypores. In: Saproxylic beetles Infrastructure. in Europe: monitoring, biology and conservation. Studia Penteriani V. and Delgado M. d. M. (2012). There is a limbo forestalia Slovenica. 137:53–58. under the moon: what social interactions tell us about Sharafia S. M., Moilanen A., White M., and Burgman M. the floaters’ underworld. Behav. Ecol. Sociobiol. (2012). Integrating environmental gap analysis with Pietola L. H., Jero J., Jalkanen R., Kinnari T., Jero O., spatial conservation prioritization: A case study from Frilander M., Pajusola K., Salminen M., and Aarnisalo Victoria, Australia. J. Environ. Manag. 112:240-251. A. (2012). Effects of p27Kip1- and p53-shRNAs on Sirkiä S. M., Lehtomäki J., Lindén H., Tomppo E., and kanamycin damaged mouse cochlea. World journal of Moilanen A. (2012). Spatial conservation prioritization otorhinolaryngology. 2:1-7. of capercaillie (Tetrao urogallus) lekking landscapes in Pouzols F. M., Barriga Barros A., R. Lopez D., and Sánchez- South-Central Finland. Wildl. Biol. 18:337-353. Solano S. (2012). Enabling Fuzzy Technologies in Sundell J., Church C., and Ovaskainen O. (2012). Spatio- High Performance Networking via an Open FPGA- temporal patterns of habitat use in voles and shrews Based Development Platform. Applied Soft Computing. modified by density, season and predators.J. Anim. Ecol.

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81:747-755. predictive adaptive response: modeling the life history of Susi H. (2012). Mustavadelman nekroosiviruksen a tropical butterfly (Bicyclus anynana). Am. Nat. tunnistaminen. Kasvinsuojelulehti. 3/2012. van Nouhuys S. (2012). Parasiternas fascinerande värld. Taberlet P., Zimmermann N. E., Englisch T., Tribsch A., exempel från åländska ängsmarker.22-24. Holderegger R., Alvarez N., Niklfeld H., Coldea G., van Nouhuys S., Niemikapee S., Hanski, I. (2012). Variation Mirek Z., Moilanen A., Ahlmer W., Marsan P. A., Bona in a host-parasitoid interaction across independent E., Bovio M., Choler P., Cieslak E., Colli L., Cristea V., populations. Insects. 3:1236-1256. Dalmas J. P., Frajman B., Garraud L., Gaudeul M., Gielly van Nouhuys S. and Kraft T. S. (2012). Indirect interaction L., Gutermann W., Jogan N., Kagalo A. A., Korbecka G., between butterfly species mediated by a shared pupal Kupfer P., Lequette B., Letz D. R., Manel S., Mansion G., parasitoid. Popul. Ecol. 54:251-260. Marhold K., Martini F., Negrini R., Nino F., Paun O., van Nouhuys S., Reudler J. H., Biere A., and Harvey J. Pellecchia M., Perico G., Piekos-Mirkowa H., Prosser F., A. (2012). Performance of secondary parasitoids on Puscas M., Ronikier M., Scheuerer M., Schneeweiss G. chemically defended and undefended hosts. Basic Appl. M., Schonswetter P., Schratt-Ehrendorfer L., Schupfer F., Ecol. 13:241-249. Selvaggi A., Steinmann K., Thiel-Egenter C., van Loo M., Van Teeffelen A., Lung T., Meller L., Vermaat J., Lavalle C., Winkler M., Wohlgemuth T., Wraber T., Gugerli F., and and Cabeza M. (2012). EU Biodiversity policy in times Consortium I. (2012). Genetic diversity in widespread of climate change. RESPONSES Policy Update. No 1. species is not congruent with species richness in alpine Van Teeffelen A. J. A., Vos C. C., and Opdam P. (2012). plant communities. Ecol. Lett. 15:1439-1448. Species in a dynamic world: Consequences of habitat Tack A. J. M., Gripenberg S., and Roslin T. (2012). network dynamics on conservation planning. Biol. Cross-kingdom interactions matter: fungal-mediated Conserv. 153:239-253. interactions structure an insect community on oak. Ecol. Virkkala R., Heikkinen, R.K., Fronzek, S., Kujala, H., and Lett. 15:177-185. Leikola, N. (2013). Does the protected area network Tack A. J. M., Johnson M. T. J., and Roslin T. (2012). Sizing preserve bird species of conservation concern in a rapidly up community genetics: it’s a matter of scale. Oikos. changing climate? Biodivers. Conserv. 22:459-482. 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 Honours and awards systems: causes and consequences. J. Evol. Biol. 25:1918- 1936. Anna-Liisa Laine received L’Oreal UNESCO for Women Thomas C. D., Anderson B. J., Moilanen A., Eigenbrod in Science award and Academy of Finland award for F., Heinemeyer A., Quaife T., Roy D. B., Gillings S., Scientific Courage. 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 Visitors P. N., Barrett L. G., and Burdon J. J. (2012). Rapid genetic change underpins antagonistic coevolution in a natural David Muru, February - September 2012. host-pathogen metapopulation. Ecol. Lett. 15:425-435. Tanya Semenova, August 2011 - February 2012. Tollenaere C., Susi H., Nokso-Koivisto J., Koskinen P., A. Dmitry Finkelstein , January 22 - 28. Tack P. A., Paulin L., Frilander M. J., Lehtonen R., and Dan Lawson, January 22 - 24. Laine A.-L. (2012). SNP Design from 454 Sequencing Cristina Banks-Leite, February 22 - 23. of Podosphaera plantaginis Transcriptome Reveals a Genetically Diverse Pathogen Metapopulation with Robert Ewers, February 22 - 23. High Levels of Mixed-Genotype Infection. PLoS One. Patrik Nosil, February 29 - March 1. 7:e52492. Greta Bocedi, March 25 - April 5. Travis J., Mustin K., Barton K., Benton T., Clobert J., Delgado Eliezer Gurarie, April 17 - May 4. M. d. M., Dytham C., Hovestadt T., Palmer S., van Dyck John Drake, May 22 - 26. H., and Bonte D. (2012). Modelling dispersal: an eco- Steve Beissinger, April 26 - May 31. evolutionary framework incorporating emigration, Brian Huntley, May 30 - June 3. movement, settlement behaviour and the multiple costs Felix Horns, August 2011 - June 2012. involved. Methods in Ecology and Evolution. Benoît Pernechele, February - June Triviño M. (2012). Global change impacts and conservation Viacheslav Spirin, April - June. priorities in the Iberian Peninsula. University Rey Juan Carlos and National Museum of Natural Sciences, Mamy Rakotoarijaona, August 24 - September 3 Madrid. Juan M. Morales, September 3 - 24 van den Heuvel J., Saastamoinen M., Zwaan B. J., Brakefield Evgen Dykyi, Oleksandr Ordynets, Mariia Pavlovska and P. M., Kirkwood T., and Shanley D. (in press). The Kseniia Tuholukova, October 7 - 10.

49 Teaching Statistics Biostatistiikka II, University of Helsinki, November Ecology - December Acroecology, Department of Agricultural Sciences, A basic course in statistics. University of Helsinki Organizer: Otso Ovaskainen Guest lecture on pathogen epidemiology and evolution in the agro-ecological interface Workshops and meetings Lecturer: Ayco Tack European boreal forest biodiversity (EBFB)- Spring Symposium 2012, Departments of Workshop on Collecting and Analyzing Winter Biosciences and Environmental Sciences, Faculty Track Data, Mekrijärvi Biological Station 11. - of Agriculture and Forestry, Museum of Natural 13.4.2012 History Members of organizing committee: Juri Kurhinen and Annual event organised by and for graduate students Otso Ovaskainen working in various fields of ecology, evolution, systematics and nature conservation. European boreal forest biodiversity (EBFB) - PhD Member of organizing committee: Tanjona Ramiadantsoa course in St Petersburg on Mathematical and Statistical Modeling of Ecological Data 23. – Ecology and Evolutionary biology weekly Seminar 27.4.2012 Series Organizers: Andrei Brodsky and Otso Ovaskainen Organizer: Anna-Liisa Laine & Heikki Hirvonen Stochastic Dynamics: Mathematical Theory and Applications reserarch program, Bielefeld, Plant-microbe interactions and molecular defence Germany, 18.5. - 14.7.2012 of plants Workshops “Stochastic Dynamics in Action” (20–25 An advanced course in molecular plant pathology. May 2012) and “Qualitative behaviour of stochastic Lecturer: Anna-Liisa Laine systems and applications” (18th June - 21th June) Member of organizing committee: Otso Ovaskainen Genetics Next Generation Genomics, University of Helsinki, Conservation biology 6 - 14 September Reserve planning in the tropics: RESPECT, Two practical courses (mapping and assembly of next Madagascar, November - December generation sequencing data; RNAseq analysis using This is a CIMO North-South-South initiative. A field R) and scientific symposium on next generation course including Malagasy and Finnish st udents, aimed sequencing methods. at teaching both ecological and development needs in Member of organizing committee: Mikko Frilander conservation planning.

Coordinators: Mar Cabeza and Jukka T. Lehtonen 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

50 MRG -ANNUAL REPORT 2012 Synopsis of the year

Conservation Biology in Fragmented Landscapes, extent to which scientific information is actually used University of Helsinki, September - November when real decisions are made. The students presented the results of these working groups during a two day Since the year 2000, the Metapopulation Research seminar in November at the Lammi Biological Station. Group has annually organized a master level course To stimulate discussion and introduce a more applied called Conservation Biology in Fragmented Landscapes perspective, we invited three external guests working (7 ECTS). The course is part of the international Boreal on applied conservation issues: Mia Valtonen from Biota and Ecology program opened to both international the University of Eastern Finland who is an expert on exchange students and the Saimaa seal, Lauri Finnish students. Our Kajander, who works objective is to unite all with forest conservation MRG members in a issues in Luonto- joint venture, to compile Liitto (the Finnish a coherent teaching unit Nature League), and that takes advantage of Juha Lumme, who our own research, and to is an expert on the provide group members flying squirrel and with essential teaching works for Uudenmaan experience. As in all our ELY-keskus (the work, we blend theory Center for economic and modelling with development, transport empirical case studies, and try to offer the students our and environment). During the seminar the students view on how habitat fragmentation affects key aspects gave two presentations per group (a detailed talk of population biology. Students are also given the choice aimed at a scientific audience and a shorter one aimed of earning two extra credits by reading books produced at a more general audience varying from land owners in the group - this year The Shrinking World by Ilkka to policy makers). The seminar was a great success, Hanski. 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

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

51 Annual meeting in St Petersburg

The annual meeting was held 3-7 October in St board also our SAB members, prof. Marian Goldsmith, Petersburg, Russia. We travelled by boat to enjoy the prof. Tony Ives and prof. Kevin Gaston, as well as the views on the archipelago and to take advantage of the Academy of Finland representative Harri Hautala. We conference facilities, thus starting the meeting already wish to thank them all once again for fruitful discussions while traveling. The program of the annual meeting and ideas! consisted of scientific presentations covering both past Concerning the social activities, we enjoyed splendid and planned work, as well as of discussions on how to meals both in Georgian and Russian restaurants. As make our group function even better than it has done evidenced by the photos, some of us celebrated our so far, both in terms of scientific and non-scientific scientific achievements all night long, while others activities. The regular annual meeting was combined focused more on cultural activities. 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

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

On a visit to the State Hermitage Museum

52 MRG -ANNUAL REPORT 2012 Synopsis of the year

Funding

Funding source EUR

Academy of Finland Centre of Excellence funding Metapopulation Research Group 1 077 974 GENERAL RESEARCH GRANTS Ilkka Hanski: The Glanville Fritillary Butterfly Genome and Population Genomics 116 190 Otso Ovaskainen: Linking Environmental Change to Biodiversity Change: Long-term and 156 236 Large-scale Data on European Boreal Forest Biodiversity Academy professor's research costs Ilkka Hanski: Eco-evolutionary spatial dynamics 281 072 Academy research fellow's research costs Anna-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 81 987 Metapopulation Dynamics Mar Cabeza Jaimejuan: Conservation efectiveness: Sociopolitical and Environmental 2 776 challenges, Academy of Finland RESEARCH INFRASTRUCTURES (FIRI 2010) Ilkka Hanski: FIRI:GenoEvo 199 500 Postdoctoral projects Anni 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 104 690 Populations Dmitry Schigel: Colonization gates and establishment of wood-decaying fungi in European 34 041 Spruce + 1 Academy professor post and 4 Academy research fellow posts

University of Helsinki Centre of Excellence funding from Ministry of Education 508 933 Ilkka Hanski: Biocentrum Helsinki 46 000 Atte Moilanen: Quantitative Methods for Targeting Conservation Management and 42 000 Intensive Forestry in Forests in Finland (three year grant) Atte Moilanen: ERC/ Moilanen (Department of Biosciences) 45 000 Atte Moilanen: ERC/ Moilanen (Bursar) 25 000 Anna-Liisa Laine: ERC/ Laine (Bursar) 25 000

53 Otso Ovaskainen: Starting Grant of the University of Helsinki Funds for Appointment as 5 000 Professor of Mathematical Ecology Atte Moilanen: Starting grant of the University of Helsinki Funds for Appointment as 6 250 Professor of Conservation Decision Analysis Ilkka Hanski: FIRI:GenoEvo (Bursar) 85 500 Swee Chong Wong: GPBM Helsinki Graduate Program in Biotechnology and Molecular 13 770 Biology, International PhD Training Program

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 10 000 and Spatial, Temporal and Ecological Scales Mar Cabeza: European Responses to Climate Change: Deep Emissions, Reductions and 212 141 Mainstreaming of Mitigation and Adaptation

OTHER FICS - FINNISH DOCTORAL PROGRAMME IN COMPUTATIONAL SCIENCES Otso Ovaskainen: Graduate school funding Sonja Koskela 2 486 EMIL AALTONEN FOUNDATION Henna Fabritius: Support grant 5 000 FINNISH FOREST AND PARK SERVICE Atte Moilanen: Support for Professorship in Conservation Decision Analysis. GIS- 60 000 based Ecological Decision Analysis in the South-Central Finland Forest Biodiversity Program (METSO) FINNISH NATURE CONSERVATION FOUNDATION Henna Fabritius: Succession of Melitaea diamina meadows 1 500 NERC UNIVERSITY OF EAST ANGLIA Otso Ovaskainen: Subcontract Agreement: The role of Dispersal in Species Ability to 7 021 Respond to Climate Change NWO NETHERLANDS ORGANISATION FOR SCIENTIFIC RESEARCH Henjo de Knegt: Rubicon grant 70 328

54 MRG -ANNUAL REPORT 2012 Synopsis of the year

OSKAR ÖFLUND FOUNDATION Silvija Budaviciute: Support grant 6 000 SOCIETAS ENTOMOLOGICA HELSINGFORSIENSIS Silvija Budaviciute: Support grant 1 000 THE CENTRES FOR ECONOMIC DEVELOPMENT, TRANSPORT AND ENVIRONMENT Henna Fabritius: Research Collaboration with South Ostrobothnia Centre, Monitoring 3 000 of the Distribution of Melitaea diamina in Western Finland Henna Fabritius: Research Collaboration with Southwest Finland Centre, Monitoring of the 2 000 Distribution of Melitaea diamina in Western Finland LUOVA-graduate school 5 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.

Other 3 %

European Academy of Union Finland 32 % 49 %

University of Helsinki 16 %

Sources of Metapopulation Research Group funding in 2012.

55 Prospects for the year 2013

ur new funding cycle as a national Centre- teaching of ecology and evolutionary biology suffering. of-Excellence (CoE) started in January Times are tough in the university, but it is inexcusable 2012, and this support will continue until that the ‘reward’ of success in raising external funding 2016.O That is the good news. The bad news is that the has a negative sign, leads to reduction of core funding. level of funding is disappointing. The support from Putting aside these very troubling prospects our host institute, the University of Helsinki, is at the concerning university funding for the year 2013, the same level as in the beginning of our previous CoE prospects for MRG are generally good. Several PhD period, in 2006, in spite of MRG now being roughly students will finish their studies in 2013, compensating twice as large, and in spite of us bringing roughly 2 for the past two years with exceptionally low numbers, million € more overheads per 6 years than in 2006. partly reflecting the shift towards us recruiting more And this is not all. Although the section of Ecology post docs rather than students a few years ago (see and Evolutionary Biology (EEB) at our department is the graph on p. 6). We didn’t manage to publish the performing really well in research and teaching, to a genome of the Glanville fritillary in 2012 as expected large extent thanks to MRG, the funding situation has – but surely this will happen in 2013! I personally look deteriorated severely also at this level. The professors forward to the next MRG dung beetle expedition, this who have left, and those whose salary is paid by the time to Gunung Mulu in Sarawak, Malaysia, where I research council, are not replaced, which means that did field work back in 1978. It is very special to go back the more successful EEB and MRG are, the more is and to have a chance to take some of you with me.

Ilkka Hanski

56 MRG -ANNUAL REPORT 2012