Metapopulation Research Group

Annual Report 2008

Department of Biological and Environmental Sciences University of Helsinki

Helsinki 2008 Contact information

Address: Metapopulation Research Group Department of Biological and Environmental Sciences 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

Printed in Picaset OY

Helsinki, December 2008 Contents

Evolution of the Metapopulation Research Group...... 5

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

Research Projects...... 9

Integrative ecology Metapopulation biology of the Glanville fritillary...... 10 Functional genomics of the Glanville fritillary...... 12 The Glanville fritillary metacommunity in the Åland islands...... 14 Metacommunity dynamics in mosaic landscapes...... 16 Metacommunity dynamics of wood-decaying fungi...... 18 Mathematical modelling Dispersal and ecological and evolutionary dynamics...... 20 Stochasticity and space in population dynamics...... 22 Large scale ecology and conservation Biodiversity informatics and conservation...... 24 Brown bear project...... 27 Biology of an old adaptive radiation: Evolutionary ecology of dung beetles in Madagascar...... 28 Other projects Ecology of small rodents and their predators...... 30 Butterfly expansion and climate change...... 31 Spatially structured coevolution...... 32 Supporting personnel...... 34

Synopsis of the year 2008...... 35 Publications...... 36 Honours, awards and memberships...... 46 Conferences and seminars...... 47 Work abroad...... 48 Visitors...... 48 Teaching...... 49 Budget...... 50 Prospects for the year 2009...... 52

Evolution of the Metapopulation Research Group

esearch groups are like local populations in a metapopulation, nodes in a vast global network. Some groups are large, others are small, partly because the amount of resources Rvary from one ‘patch’ (institute) to another, partly because of variation in the many other factors affecting the sizes and dynamics of research groups. New groups are constantly being established while others gradually dwindle to extinction. There is dispersal and gene flow among existing research groups.

Research groups are typically established around a single researcher. Successful groups then increase in size – in the numbers of students, post docs, and technicians. In 5-10 years time, the growth becomes limited by lack of resources such as space and funding but also by the time that the senior researcher has available for running the group. At this point a successful group starts sending dispersers to other groups and even to establish new groups elsewhere - to compete for resources away from the natal patch.

All this has happened in the case of the Metapopulation Research Group (MRG), but our group has additionally undergone a transformation from a research group of a single researcher to a large group jointly run by several researchers. There is now important internal structure within MRG, each senior researcher having his or her own students and post docs and partly own funding.

MRG has been retained as an umbrella organization partly for funding purposes – we have been competitive as a group for the Centre-of-Excellence funding from the Academy of Finland (national research council). But there is more to it. MRG has evolved into a large group with a range of interests and expertises from mathematics and computer science to ecology, population biology, and conservation biology. In the past few years, we have started to move into molecular biology, genomics, and bioinformatics. This latter expansion is still happening, but it is already apparent that in a few years time MRG will be quite a beast.

Evolution of research groups is an interesting and important phenomenon, not least because it will set the stage for the type of academic training and research that is going on. There is no single path that every research group should follow, diversity of groups adds to the strength of the network of groups as a whole. We have become a large group in terms of both human and material resources. We should aim to make contributions to research that are not possible without these resources. These contributions include further integration of theoretical and empirical research in population biology, which has been our strength in the past, but also meaningful integration of molecular- level and population-level studies.

Ilkka Hanski

©Tapio Vanhatalo

5 Brief history and overview of MRG

etapopulation Research Group was Current status established by Ilkka Hanski in 1991. MHanski had worked on spatially structured MRG is the leading research group worldwide in populations since the late 1970’s. The early metapopulation biology and one of the Centres-of- work dealt with small-scale spatial structure Excellence in Research nominated by the Academy within populations, but since the early 1980’s of Finland (national research council) in 2006-11. the focus shifted to larger spatial scales and to Our past strengths include successful integration metapopulation dynamics. Important events leading of theory, modelling, and empirical research, as to the establishment of MRG include the first well as the development of the Glanville fritillary international meeting on metapopulation dynamics butterfly into a widely recognized model system in organized by Hanski and Michael Gilpin (San Diego, metapopulation biology. The senior researchers who US), which resulted in the first edited volume on have joined MRG as group leaders in the past years the subject (Gilpin & Hanski, 1991, Metapopulation have opened up new fronts of research in bringing Dynamics: Empirical and Theoretical Investigations, together metapopulation biology and state-of- Academic Press, London). The long-term and the-art mathematical modelling (Ovaskainen) large-scale metapopulation project on the Glanville and functional genomics (Frilander) as well as fritillary butterfly (Melitaea cinxia) in the Åland incorporating spatial dynamics into research on the Islands in SW Finland was started in 1991. design of reserve networks (Moilanen).

Current structure General goals for research The graph shows the growth of MRG since 1992. Our vision of top-quality interdisciplinary research At present, MRG is a highly international group of in population biology and evolutionary biology 40 senior researchers, post docs, post graduate involves work that is motivated by biological students, and supporting personnel representing questions but which at the same time may make an 11 nationalities. The senior researchers are leaders original contribution to basic research in the other of their own groups of students and post docs as disciplines that are represented in MRG: molecular described on p. 7. The senior researchers represent a biology, mathematics, and computer science. The wide range of expertises, including ecology (Hanski, work includes both basic research and applications Roslin, van Nouhuys), computer science (Moilanen), to conservation and management. Our CoE position conservation biology (Cabeza), mathematics has allowed us to initiate projects cutting across (Ovaskainen) and molecular biology (Frilander). interdisciplinary boundaries and to build the

Fig. 1. MRG personnel since 1992.

6 infrastructure needed for future work. The three research, spanning from basic mathematical work general goals of research in the period 2006-11 are: on new theory to statistically rigorous modelling of particular systems. 1. Integration of molecular, population and evolutionary biology in the spatial context 3. Development of new approaches in large- scale ecology and conservation in the context of Previous research in MRG was largely focused on ecological spatial dynamics. A fundamental global changes understanding of many biological processes calls Previous research in metapopulation ecology for integration of molecular biology with ecology has been primarily focused on processes at the and evolutionary biology. We have designed a new landscape level, whereas the major drivers of research program to achieve this combination in the global change often operate at very large scales. context of our long-term project on the Glanville We have established a comprehensive research fritillary butterfly. program on the design of reserve networks, which will address the influence of long-term habitat 2. Development of mathematical theory and change and climate change. The planned work predictive models for spatial dynamics includes the development of new methods as well as their applications. The methodological work Previous research in MRG has involved the incorporates habitat modelling, spatial optimisation, construction of metapopulation models that can and uncertainty analysis into advanced large-scale be parameterised for real metapopulations. The computational tools for reserve planning. mathematical and computing competence of the current researchers and students in MRG allows us to set significantly more ambitious goals for

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

Senior researcher Ilkka Hanski Senior researcher Otso Ovaskainen Phil Harrison 1 Eliezer Gurarie Ines Klemme Juho Pennanen Chris Wheat 2 Jenni Hottola3 Varpu Mitikka Veera Norros Kristjan Niitepõld Ace North Pierre Rahagalala Chaozhi Zheng Heidi Viljanen Helena Wirta Senior researcher Tomas Roslin Riikka Kaartinen Senior researcher Mar Cabeza Ayco Tack Heini Kujala Independent Senior researcher Mikko Frilander researchers Saskya van Nouhuys Jouni Kvist3 Olivier Gilg Elina Niemelä Anna-Liisa Laine Heli Pessa Janne Sundell Janne Turunen Jens Verbeeren 1 in collaboration with OO Senior researcher Atte Moilanen 2 in collaboration with MF Anni Arponen 3 co-supervised by IH Joona Lehtomäki3

Research Projects

Integrative ecology Metapopulation Biology of the Glanville fritillary Functional genomics of the Glanville fritillary The Glanville fritillary metacommunity in the Åland islands Metacommunity dynamics in mosaic landscapes Metacommunity dynamics of wood-decaying fungi

Mathematical modelling Dispersal and ecological and evolutionary dynamics Stochasticity and space in population dynamics

Large scale ecology and conservation Biodiversity informatics and conservation Brown bear project Biology of an old adaptive radiation: Evolutionary ecology of dung beetles in Madagascar

Other projects Ecology of small rodents and their predators Butterfly expansion and climate change Spatially structured coevolution

Supporting personnel

Ilkka Hanski Kristjan Niitepõld professor PhD -student Phil Harrison Luisa Orsini post doc post doc

Undergraduate students: Eveliina Kallioniemi, Anniina Mattila

he large metapopulation of the Glanville of ca 4,000 habitat patches. The size of the fritillary butterfly (Melitaea cinxia) in the Åland metapopulation decreased in 2007 due to TIslands in SW Finland has been studied since periods of dry weather in summer increasing 1991. Over the years, this project has expanded larval mortality. from a case study of metapopulation ecology to a widely recognized model system in population • Individuals with a particular variant of the gene biology, with many important contributions to the phosphoglucose isomerase (Pgi) have been study of the ecological, genetic, and evolutionary shown to have superior dispersal capacity and consequences of habitat fragmentation. The fecundity in the Glanville fritillary butterfly, Glanville fritillary project has also provided an raising questions about the mechanisms that example of how theoretical and empirical research can be effectively combined. This project is closely linked with three other projects in MRG, on metacommunities, spatial modeling, and functional genomics. The functional genomics studies will expand greatly in the coming years, supported by a big grant from the European Research Council. The aim of this 5-yr project is to advance our general understanding of the genetic basis of variation in individual performance and life-time reproductive success (fitness), and of the role of ongoing natural selection in population dynamics of species living in fragmented landscapes.

Highlights of the research in 2008: Fig. 1. Changes in the numbers of larval families in the large metapopulation of the Glanville fritillary in • The long-term observational study of the the Åland Islands in SW Finland since 1993. Note that metapopulation in the Åland Islands has there has been no change in the overall size of the been continued by surveying the presence of metapopulation, but the amplitude of fluctuations local populations in April (n=707) and again appears to have increased, perhaps due to more in the autumn (n=513) in the large network frequent unusual weather events.

10 maintain polymorphism in this gene in natural in flight muscles and consequently enrichment populations. In an experiment run at the of certain splice forms due to selective loss of Lammi Biological Station, Marjo Saastamoinen, muscle cells. Suvi Ikonen and Ilkka Hanski investigated how variation in the Pgi genotype affects • Luisa Orsini and colleagues have compared the female and male life history under controlled spatial genetic and demographic structures conditions. The most striking effect was longer in the Glanville fritillary metapopulation in lifespan of genotypes with high dispersal the Åland Islands. The results indicate that capacity. Butterflies used body reserves for the spatial genetic structure in a particular somatic maintenance and reproduction, but year corresponds better to the demographic different resources (in thorax vs abdomen) were structure some 5 years previously rather than used under dissimilar conditions, with some to the concurrent demographic structure. Thus interactions with the Pgi genotype. the genetic spatial structure across a large area reflects partly past demographic events. • A collaborative study with Prof. James Marden (Penn State) on the alternatively spliced forms of the flight muscle protein troponin-t in the Glanville fritillary points to a possible mechanism that may allow butterflies to retain adequate flight capacity in spite of loss of flight muscle mass in the course of adult life. The relative frequency of the large Tnt isoform, which is expected to promote force and power output, increased with decreasing ratio of thorax mass at the end of life to initial body mass. This result suggests that there may be non-random distribution of the Tnt isoforms

Collaborators

s 2.2 Prof. James Marden, Penn State University, USA in Prof. Rongjiang Wang, Peking University, China 2.1 5 m

in 2.0 Key publications

ed 1.9 Saastamoinen, M. and Hanski, I. (2008). Genotypic ov 1.8 and environmental effects on flight activity and

e m 1.7 oviposition in the Glanville fritillary. The American Naturalist 171, E701-712.

tanc 1.6 Orsini, L., Corander, J. Alasentie, A. and Hanski, I. is Pgi AA (2008). Genetic spatial structure in a butterfly 1.5 Pgi AC g d metapopulation correlates better with past than

Lo 1.4 present demographic structure. Molecular Ecology 14 16 18 20 22 24 26 28 30 17, 2629-2642. Marden, J.H., Fescemyer, H.W., Saastamoinen, M., Ambient temperature °C MacFarland, S.P., Vera, J.C., Frilander, M.J. and Fig. 2. Harmonic radar-measured dispersal rate in the Hanski, I. (2008). Weight and nutrition affect pre- Glanville fritillary in relation to Pgi genotype and ambient mRNA splicing of a muscle gene associated with air temperature. The dots give the model-predicted performance, energetics and life history. Journal dispersal rate. Pgi AC heterozygotes (SNP AA111; red dots) of Experimental Biology, 211, 3653-3660. moved actively already in low ambient temperatures Orsini, L., Wheat, C.W., Haag, C.R., Kvist, J., Frilander, whereas AA homozygotes (blue dots) moved longer M.J. and Hanski, I. (in press). Fitness differences distances only in higher temperatures. The difference associated with Pgi SNP genotypes in the between the genotypes may be due to dissimilar effects of Glanville fritillary butterfly (Melitaea cinxia). temperature on enzyme kinetics of the two isoforms of the Journal of Evolutionary Biology. PGI enzyme (Niitepõld et al. Ecology in press).

11 Functional genomics of the Glanville fritillary

Mikko Frilander Chris Wheat Ilkka Hanski group leader post doc professor

PhD -students in RNA-splicing project: Elina Niemelä, Heli Pessa, Janne Turunen, Jens Verbeeren

Technicians: Marja-Leena Luisa Orsini Jouni Kvist Peltonen, Annukka Ruokolainen post doc PhD -student

olecular level investigations with Glanville fritillary focus on studying the effect of Mgenetic variation on dispersal rate and other life history traits in the Åland metapopulation. We investigate genetic variation at three different levels, namely transcriptional variation, allelic variation, and variation at the level of alternative pre-mRNA splicing. Currently we are using microarrays to study transcriptional variation. With allelic variation our focus is on phosphoglucose isomerase (Pgi), and we are investigating variation in biochemical performance among its amino acid variants using a variety of biochemical and molecular biology methods. Our candidate gene for the alternatively splicing variation is the muscle protein gene troponin T.

Highlights of recent work are: Fig. 1. Venn diagram showing the number of genes 1. Sequencing of the Glanville fritillary that have a significant expression difference (p-value transcriptome. We used massive parallel < 0.05) in developing last instar larvae of the Glanville sequencing (454 sequencing, Roche Ltd.) to fritillary butterfly (Melitaea cinxia), contributed to identify more than 50% of the coding region growth temperature, larval family or the interaction of of M. cinxia genome. This was the firstde novo temperature and family. Expected false discovery rate in transcriptome sequencing with massive parallel parenthesis.

12 technology ever done for any multicellular organism (Wheat, Hanski, Frilander; Marden, Fescemyer, Vera at Penn State University; Crawford at University of Miami). Fig. 3.Cover of Molecu- 2. Sequence data was used to construct Agilent lar Ecology (Vol 17, issue oligonucleotide microarrays. Two microarray 7) depicting the benefits experiments are in progress: the influence of of M. cinxia transcrip- growth temperature and genetic background tome: Discovery of a large on larval gene expression (Kvist, Wheat, Hanski, number of SNP alleles and Frilander; Marden at Penn State), the influence development of an oligo- of metapopulation structure on adult gene nucletide microarray for expression (Wheat, Kvist, Hanski, Frilander; gene expression studies. Marden at Penn State University).

3. We have uncovered a connection between Collaborators pre-mRNA splicing of the Troponin T and life-history variables (nutritional status, Prof. James Marden, Penn State University, USA metabolic performance) in M. cinxia (Frilander, Prof. Reinhard Lührmann, Mac Planck Institute, Saastamoinen, Hanski; Marden, Fesmayer, Vera Göttingen, Germany MacFarland at Penn State University). Prof. Olli Silvennoinen, Institute of Medical Technology, University of Tampere, Finland 4. We are investigating single nucleotide polymorphism (SNP) distribution in M. cinxia Key publications populations in Åland, France, and China using Vera J.C., Wheat C.W., Fescemyer H.W., Frilander 454 FLX sequencing methods (Kvist, Wheat, M.J.,Crawford D.L., Hanski I. & Marden J.H. (2008). Frilander, Hanski; Marden, Vera at Penn State Rapid transcriptome characterization for a non- University). model organism using massively parallel 454 pyrosequencing. Molecular ecology, 17, 1636- 1647. Marden J.H., Fescemyer H.W., Saastamoinen M., MacFarland S.P., Vera J.C., Frilander M.J., & Hanski I. (2008). Weight and nutrition affect pre-mRNA splicing associated with energetics and life history. Journal of Experimental Biology, 211, 3653-3660. Orsini L., Wheat C.W., Haag C.R., Kvist J., Frilander M.J. & Hanski, I. (in press). Fitness differences associated with Pgi SNP genotypes in the Glanville fritillary butterfly (Melitaea cinxia). Journal of Evolutionary Biology.

Key Publications for mRNA splicing project

Pessa, H.K.J., Will, C.L., Meng, X., Schneider, C., Watkins, N.J., Perälä, N., Nymark, M., Turunen, J.J., Lührmann, R., and Frilander, M.J. (2008). Minor spliceosome components are predominantly localized in the nucleus. Proceedings of the National Academy of Sciences USA 105, 8655- 8660. Turunen, J.J., Will, C.L., Grote, M., Lührmann, R., Fig. 2. Heatmaps of the significant genes for temperature and Frilander, M.J. (2008). The U11-48K Protein (Cold, Standard & Hot) and family (N170, N74 & O171). Contacts the 5’ Splice Site of U12-Type Introns Hierarchical clustering is based on the standardized and the U11-59K Protein. Molecular and Cellular residuals of the mean expression difference from a mixed Biology 28, 3548-3560. model analysis.

13 The Glanville fritillary metacommunity in the Åland islands

Undergraduate students: Tom Kraft1, Markku Ojanen2

1 (Cornell University) 2 (co-supervised by Anna-Liisa Laine) Saskya van Nouhuys senior researcher

his branch of the metacommunity project narrow host ranges. This summer we did a large started in the early 90s with the study of the field experiment testing for apparent competition Tspatial dynamics of two primary parasitoids of between the butterflies due to the two generalist the Glanville fritillary butterfly, Cotesia melitaearum parasitoids (fig 1). We found that parasitism of and Hyposoter horticola. Gradually we have M. cinxia was decreased by the presence of M. broadened our study to include other parasitoids, athalia (fig. 2). This pattern was evident at the the host food plant, a plant pathogen, and a scale of our 5 m2 replicated plots, suggesting that related butterfly, Melitaea athalia. Our current work at least at a local scale M. cinxia benefits from the falls into the general categories: metacommunity co-occurrence of M. athalia. At a regional scale, ecology, population dynamics, and behaviour. parasitism of both butterflies was low in plots located in areas of low butterfly density, and high Metacommunity ecology. In 2008 we made where the butterfly density was high. progress toward understanding the indirect interaction between butterfly species, mediated by Population dynamics. This autumn we also began their shared parasitoids. The butterflies Melitaea a thorough study of the spatial distribution of cinxia and M. athalia share four pupal parasitoid larval parasitoids and hyperparasitoids of M. cinxia species, two with wide host ranges and two with in conjunction with other work on the genetics

Fig. 1. Tom Kraft sitting next to the two species treatment plot of the pupal parasitism experiment. Mesh covered potted Veronica spicata plants each contain one last instar Melitaea cinxia or M. athalia larva.

14 Fig. 3. Andy Philippides observing as a video camera records Hyposoter horticola making learning flights Fig. 2. The fraction of Melitaea cinxia and M. athalia around a central Veronica spicata plant with host eggs on pupae parasitized by Pteromalus apum and P. puparum it, surrounded by plants without host eggs on them. in low density plots (single species) and high density plots (both species in a “checkerboard” pattern). Each treatment is replicated 11 times, each replicate in a different part of the Åland islands. and metapopulation biology of the butterfly. We 3). We will use complex analysis of video imaging obtained samples of parasitoids from multiple to compare the details of the learning flights of H. host butterfly families from each local butterfly horticola with those of social honeybees and solitary population in Åland. This allows us to clearly bumblebees. Our expectation is that though the associate parasitoid population dynamics and purposes of the learning flights differ (foraging site metacommunity structure with host population vs. nest site), these species are phylogenetcially connectivity, age and size. related, so their behaviour may have common components. In collaboration with A.L. Laine we published a study showing the surprising positive association Collaborators of parasitoid metapopulation dynamics with the occurrence of a plant pathogenic fungus (van Prof. Tom S. Collett, University of Sussex, UK Nouhuys & Laine 2008). This is partially based Dr. Andy Philippides, University of Sussex, UK on laboratory work conducted by gradu student Prof. Donald Quicke, Silwood Park, UK Markku Ojanen, showing that C. melitaearum Dr. Joanneke Reudler Talsma, University of Jyväskylä parasitoids developing in host caterpillars feeding Dr. Mark Shaw, National Museum of Scotland on fungus infected plants tend to be female. The large-scale positive association may be because persistence of newly colonized populations of the Key publications parasitoid increases with a female biased sex ratio. Reudler-Talsma, J., Torri, K. van Nouhuys, S. (2008). Parasitoid foraging behaviour. Over the last few Host plant use by the Heath fritillary butterfly, years we have been studying spatial learning by Melitaea athalia: plant habitat, species and the parasitoid wasp H. horticola (see van Nouhuys chemistry. Arthropod-Plant Interactions, 2, 63-75. & Kaartinen 2008). This wasp uses spatial learning van Nouhuys, S. Kaartinen, R. (2008). A parasitoid to take advantage of a host that is present in the wasp uses landmarks while monitoring potential environment for a weeks, but only available for resources. Proceedings of the Royal Society B, 275, parasitism for an extremely short interval of time. 377-385. This year, in collaboration with T. S. Collett and van Nouhuys, S. and Laine, A-L. (2008). Population A. Philippides we began a study comparing the dynamics and sex ratio of a parasitoid altered by behaviour of H. horticola with other Hymenoptera fungal infected diet of host butterfly. Proceedings that have well characterized spatial learning (fig. of the Royal Society B, 275, 787-795.

15 Metacommunity dynamics in mosaic landscapes

Tomas Roslin senior researcher Riikka Kaartinen PhD -student Ayco Tack PhD -student Undergraduate students: Bess Hardwick, Juha Höykinpuro

n this part of the metacommunity project, extend between years, experimental treatments we focus on two model systems inhabiting were established in 2007 and results collected in Ipatchy habitats: dung beetles on pastures, 2008. The results will form an integral part of PhD and specialist herbivores of the oak Quercus student Ayco Tack’s exploration of how different robur. By working on two distinct systems, we forms of competition contribute to shape local hope to gain more general insight into how communities of oak-specific herbivores. spatial context affects the interplay between individuals, populations and communities. In 2008, One of our most exciting results this year emerged we implemented one ambitious project in each from the dung beetle system. To examine how system: current habitat fragmentation may ultimately affect an important In the dung beetle system, we used volunteers ecosystem function – the from the Finnish 4H-Federation to sample 134 decomposition of dung cattle farms throughout the country (Fig. 1). – we used a microcosm This project offers a direct follow-up of a similar experiment (Fig. 3). Here, sampling conducted in 1996 (cf. Roslin 2001. we took advantage of a Ecography 24: 511-524). By re-sampling the previous, observational original farms (or their nearest neighbours) by study (Roslin & Koivunen original techniques, we expect to gain rare insight 2001, Oecologia 127: into recent changes in the Finnish dung beetle 69-77) to create realistic fauna, and into the nation-wide metacommunity differences in dung beetle dynamics of a functionally important taxon. assemblage structure representative of a In the oak system, we completed a two-year fragmented and an experiment on apparent competition (Fig. 2). Here, intact landscape. PhD student Riikka Kaartinen has built quantitative We observed that food webs of the leaf-mining and galling insect different species communities. We now use these webs to understand how different herbivore species are linked to Fig. 1. A map of cattle each other through shared natural enemies, and farms sampled by test our understanding by an empirical experiment. volunteers in 2008 Since we are primarily interested in how effects

16 contribute unevenly to the function measured, with much of dung removal depending on a single species (Fig. 4). Hence, our findings suggest that the functional outcome of habitat loss will depend on taxon-specific responses to landscape modification. Only by addressing these responses may we predict the actual consequences of habitat loss.

Fig. 3. Our microcosm experiment aimed at identifying the impact of habitat fragmentation on the decomposition of cattle dung.

© Tomas Roslin Fig. 4. Dung weight after 49 days in the presence or absence of Geotrupes stercorarius. The addition Fig. 2.Oaks used in our experiment on apparent or removal of other taxa had much smaller effects competition. Colourful flags identify what species of moth (cf. Rosenlew and Roslin 2008). was introduced where.

Collaborators Ecology 77, 757-767. Gripenberg, S. & Roslin, T. (2008). Neither the devil Prof. Pertti Pulkkinen, Finnish Forest Research nor the deep blue sea: larval mortality factors Institute, Finland fail to explain the abundance and distribution of Prof. Jens Roland, University of Alberta, Canada Tischeria ekebladella. Ecological Entomology 33, Dr. Juha-Pekka Salminen, University of Turku, Finland 346-356. Dr. Graham Stone, University of Edinburgh, UK Rosenlew. H. and Roslin, T. (2008). Habitat fragmentation and the functional efficiency of Key publications temperate dung beetles. Oikos 117, 1659-1666. Roslin, T. and Salminen, J. P. (2008). Specialization Gripenberg, S., Ovaskainen, O., Morriën, E. and pays off: contrasting effects of two types of Roslin, T. (2008). Spatial population structure of tannins on oak specialist and generalist moth a specialist leaf-mining moth. Journal of Animal species. Oikos 117, 1560-1568.

17 Metacommunity dynamics of wood-decaying fungi

Otso Ovaskainen senior researcher

Jenni Hottola PhD -student

Veera Norros Undergraduate students: PhD -student Elina Karhu, Inka Silfsten

his project is focused on metacommunity is difficult to study directly. This year we tested the dynamics of wood-decaying fungi living in possibility of using DNA-based methods to identify Tthe dynamic habitat provided by dead trees. the species from saw-dust samples (Figure 2). Polyporous fungi are among the most important wood-decaying species in boreal forests. Thirty-seven The field season was busy especially due to per cent of polypores in Finland are red-listed, mainly dispersal related experiments. In her MSc study, due to large-scale reduction in the area of natural Elina examined the reduction in viability the spores forests and the amount of dead wood in all forests. experience due to exposure to the harsh physical Our aim is to understand why some species have declined more than others, and what kind of management measures would be most effective in improving the viability of wood- decaying fungi in managed forests.

Jenni’s PhD work is based on fruit body inventories, with the aim of examining how the amount, quality and spatiotemporal availability of dead wood affects the dynamics of polypore communities and especially the occurrence of red-listed species. The aim of Veera’s PhD project is to gain a mechanistic understanding of the determinants of dispersal in wood-decaying fungi and to pin-point the factors that create differences between species in their dispersal ability. We have become increasingly Fig. 1. We used a synthetic aerosol as a model for spore dispersal. In this interested in the interactive processes experiment we measured under forest conditions the dispersal kernel for that take place inside the logs. particles in the size range relevant for fungi spores (aerodynamic diameter Unfortunately this stage of the life-cycle 1-10 µm).

18 Fig. 2. Results from a pilot study testing whether it is possible to identify wood 1.0 0.8 1.0 0.99 0.95 decaying fungi from environmental 0.4 0.99 0.7 samples. We extracted DNA from saw-dust 0.4 0.95 samples originating from spruce logs, run 0.8 0.8 the material through 454-pyrosequecning, and identified the species using a reference 0.975 library for popypores and corticoids. 0.5 0.975 0.5 In the figure each colour represents a 0.6 0.6 candidate species returned by the BLAST 0.1 0.8 0.7 0.1 alogorithm, the size of the circle being proportional to the number samples. The 0.4 0.4 contour lines depict the probabilities that the identification is correct to the species 0.3 0.9 0.3 0.9 level, estimated with the help of replicate sequences of known species. The BLAST 0.6 0.2 0.2 0.6 results with a low similarity score are likely to originate from species that are missing Uniqueness of the best BLASTUniqueness of from the reference library. The BLAST results with a low uniqueness score are relate to 0.2 0.0 0.0 0.2 samples that are not possible to relate 0.0 0.2 0.4 0.6 0.8 1.0 0.0 0.2 0.4 0.6 0.8 1.0 to the species level due to little variation between species in some of the families. Similarity score of the best BLAST

conditions of the atmosphere. She found that the germination of all seventeen measured polypore species was impeded by intensive solar radiation, Collaborators while the effect of freezing was less uniform across the species. Inka’s MSc work assessed the extent Prof. Timo Vesala, Department of Physical Sciences, to which polypore-associated beetle species fulfill University of Helsinki the preconditions to function as vectors for spores. Dr. Timo Mäkelä, Finnish Meteorological Institute She collected beetle specimen from fruiting bodies Dr. Raisa Mäkipää, Dr. Reijo Penttilä, and M. Sc. Juha to observe how well spores become attached and Siitonen, Finnish Forest Research Institute detached to different species and, with the help of M. Sc. Dmitry Schigel and M. Sc. Otto Miettinen, literature and experts, identified beetle species with a Finnish Museum of Natural History, University of life history enabling spore transport. Helsinki

To examine the effect of particle size on aerial transport, we studied the spread of a synthetic Key publications aerosol with particle size distribution covering the Hottola, J. & Siitonen, J. (2008). Significance of spore size range of wood-decaying fungi. The decay woodland key habitats for polypore diversity and of particle concentration with distance was measured red-listed species in boreal forests. Biodiversity using aerosol instruments, and measurements were and Conservation 16, 2559-2577. made both inside a forest and in an open area to Ovaskainen, O., Nokso-Koivisto, J., Hottola, J., Rajala, observe the effect of the canopy (Figure 1). Also, we T., Pennanen, T., Mäkipää, R., Ali-Kovero, H., conducted an extensive experiment comparing the Miettinen, O., Oinonen, P., Larsson, K-H, Paulin, L. dispersal kernels of three wood-decaying species and Auvinen, P. (Manuscript). Pyrosequencing as with contrasting spore size and shape. Here, fruiting a tool for metagenomics: is it possible to identify bodies of the study species were transported to a wood-decaying fungi to the species level? study area of homogeneous forest, and the decrease in spore deposition with distance was measured using species-specific spore traps.

19 Dispersal and ecological and evolutionary dynamics

Otso Ovaskainen Ilkka Hanski senior researcher Juho Pennanen Elizabeth Crone professor post doc visiting researcher

Eliezer Gurarie Chaozhi Zheng post doc Phil Harrison PhD -student post doc

his project focuses on integrating data and framework features such as movement corridors, theory in spatial ecology and evolutionary barriers, and autocorrelated movements. Eli Tbiology. One uniting aspect of the various focused on continuous-time random walks, subprojects is the development of Bayesian state- developing both mathematical aspects and space modelling approaches that combine a state applying the models to data on marine mammals process modelling the true but unknown state and other organisms. of the population, and an observation process linking the state process to a time series of With respect to spatial population processes, the observational data. modelling work continues to be closely related to the Glanville fritillary project (page 10). Phil Last year we completed the development of has developed an individual-based model that a diffusion-based modelling framework that integrates census data, mark-recapture data, and can be used to extract movement parameters life-history data. This year he worked on the from mark-recapture data (Ovaskainen et al. Bayesian parameterization of the model with 2008). This year we started the extension of this an emphasis on accounting for observation framework towards more general process models errors. Chaozhi constructed a model of dispersal (e.g. home-range movements of mammals) evolution, examining the feedbacks between and observation models (e.g. GPS data). These ecological dynamics and evolutionary dynamics developments were carried out by our Fullbright in the PGI gene that affects the metabolic visitor Elizabeth, and two new post docs, Juho performance of the Glanville fritillaries (Zheng et and Eli. Juho implemented into the diffusion al, ms). Thanks to the extensive genetic sampling

20 Theor Ecol (2008) 1:29–43 37

Fig. 6 Two examples 10 10 of restricted spatial distributions ab of equal competing species. Different colors indicate the 8 8 presence of different species with pix>0.2. a Very strong competition, β=0.9, and 6 6 b moderately strong competi- Y Y tion, β=0.7. Values of other 4 4 parameters: p� 0:8 and α=3 s ¼

2 2

0 0

0246810 0246810 Theor Ecol (2008) 1:29–43 37 X X Fig. 6 Two examples 10 10 of restricted spatial distributions ab of equal competing species. In the case of interspecific variation in ps� or β, the results with intermediate α went extinct (1.34, 2.14, 2.41, 2.51). Different colors indicate the 8 8 presence of different species are straightforward: The species that persist are always the Considering the spatial distributions of the six coexisting with pix>0.2. a Very strong ones with the largest values of ps� (best colonizers) or the species, the pattern was largely nested (Fig. 7a). The parts competition, β=0.9, and 6 best competitors (largest β values6 in model A, smallest of the patch network in which the less common species had b moderately strong competi- Y values in model B). The actual numberY of species coexisting high incidences were the ones with the highest level of tion, β=0.7. Values of other 4 at equilibrium depends on the values4 of the two other landscape connectivity (Fig. 7b). parameters: p� 0:8 and α=3 s ¼ parameters, with no variation among the species. A well-established hypothesis about competitive coexis-

2 In the case of interspecific variation2 in α and so strong tence in patchy habitat posits a trade-off between compe- competition that only one or a few species persisted, the tition and colonization (Stone 1995; Tilman et al. 1994, species that persisted were the ones with the largest values 1997; for a recent empirical example see Cadotte et al. 0 0 of α and, hence, the most local migration. When compe- 2006). In the present context, this can be interpreted as an 0246810 0246810 tition was somewhat less strong and, consequently, more interaction between ps� and β in affecting coexistence. To species coexisted,X the persisting species typically includedX test this, I generated communities of initially 10 species

also some species withFig. small 2.α Two(long-range examples migration), of restricted spatialwith the distributions values of p s� and β drawn independently for each Fig. 1. Comparison between predictedwhereas (a) and observed species with intermediate α were excluded. species from the uniform distributions in Table 1. I repeated In the case of interspecific variation in p or β, the results with intermediateof competingα went species, extinct different (1.34, 2.14,colors 2.41,indicating 2.51). the (b) allelic variation in the PGI gene s�amongFigure sub-networks7 gives an example in which 6 out of the 10 initial the calculations for α=1 and 3 and for models A and B. are straightforward: The species that persist are always the Consideringpresences the spatial of different distributions species. of The the snapshots six coexisting shown in the inhabited by the Glanville fritillary. Bothspecies panels coexisted. show the The twotwo common panels speciesare based had on the assuming largest eitherIn (a) model very A,strong there or was no trade-off: The species that ones with the largest values of ps� (best colonizers) or the species, the pattern was largely nested (Fig. 7a). The parts frequency of the C allele (associated to valueshigh mobility) of α (3.59, 3.90), whereas the four other species persisted were always the ones with the largest values of ps� best competitors (largest β values in model A, smallest of the patch(b) moderately network in whichstrong the interspecific less common competition. species had in sub-networks of habitat patches as aincluded function species of the with the next largest α (3.26, 3.35) and the (best colonizers). This is not unexpected because the values in model B). The actual number of species coexisting high incidences were the ones with the highest level of pooled number of larval groups in the networkvery smallest at theα time(0.03, 1.05). The remaining four species competitive effects experienced by species in a multispecies at equilibrium depends on the values of the two other landscapeCollaborators connectivity (Fig. 7b). parameters,of sampling with no Panel variation (a) shows among a theprediction species. of a simulation A well-established hypothesis about competitive coexis- model, panel (b) an empirical result. In the regression lines In the case of interspecific variation in α and so strong tence inProf. patchy Juha habitat Merilä posits & Dr. a trade-off Cano Arias, between Univ. compe- of Helsinki the networks in which the C allele was absentFig. 7 a (frequencySpatial distributions competition that only one or a few species persisted,of two common the andtition four less andProf. colonization Elja Arjas, (Stone Department1995; Tilman of Mathematics et al. 1994, and 0) have been excluded. species that persisted were the ones with the largestcommon values species with1997 unequal; forαStatistics, a recent empirical University example of Helsinki see Cadotte et al. of α and, hence, the most local migration. Whenvalues. compe- Different colors2006indicate). In the present context, this can be interpreted as an the presence of species with Dr. Mikko Kuussaari, Finnish Environment Institute tition was somewhat less strong and, consequently, more interaction between p� and β in affecting coexistence. To pix>0.3. The default landscapeProf. Miska Luoto,s University of Oulu speciesof coexisted,the empirical the persisting system, species we were typically ablein Table included to1 testwas assumed.thetest this, I generated communities of initially 10 species also somemodel species predictions with small againstα (long-range independentParameter migration), data values: p�with0:9 the, values of p and β drawn independently for each s ¼ s� whereas(Figure species 1). with intermediate α wereβ=0.7, excluded. and α as explainedspecies in from theKey thepublications uniform distributions in Table 1. I repeated text. b Landscape connectivity Figure 7 gives an example in which 6 out of theof the 10 patches initial in thethe network calculations in for α=1 and 3 and for models A and B. speciesIlkka coexisted. extended The two single-species common species metapopulation hada, calculated the largest for patch iInas modelOvaskainen, A, there O., was Rekola, no trade-off: H., Meyke, The species E. and thatArjas, e 2dij A0:5, where d is the valuesmodels of α (3.59, into 3.90),metacommunity whereas the fourmodels other� by species jassumingpersistedij wereE (2008). always Bayesian the ones with methods the largest for valuesanalyzing of ps� j i includedthat species interspecific with the next competition largest α (3.26, reduces 3.35)distanceP6¼ and the between the local patches(best i and colonizers).j.movements This in is heterogeneous not unexpected landscapes because the from very smallestcarryingα capacity.(0.03, 1.05). He Thefound remaining that short-distanceThe four area species of the dot incompetitive this figuremark-recapture effects experienced data. by speciesEcology in a89, multispecies 542-554. is proportional to the Zheng, C., Ovaskainen, O. and Hanski, I. (Manuscript) dispersal combined with strong competitionconnectivity of theleads to spatial pattern formation (Figurerespective 2), decreasing habitat patch Modelling single nucleotide effects in the Pgi Fig. 7local a Spatial but distributions increasing global species diversity. gene on dispersal in the Glanville fritillary of two common and four less butterfly: coupling of ecological and evolutionary commonOtso species approached with unequal α the same question from an values.empirical Different colors perspective,indicate developing multivariate dynamics. the presencelogistic of species regression with models for inferring species Hanski I. (2008) Spatial patterns of coexistence of pix>0.3. The default landscape competing species in patchy habitats. Theoretical in Tableinteractions1 was assumed. from census data. Ecology 1, 29-43. Parameter values: ps� 0:9, β=0.7, and α as explained¼ in the text. b Landscape connectivity of the patches in the network in 21 a, calculated for patch i as 2dij 0:5 e� Aj , where dij is the j i distanceP6¼ between patches i and j. The area of the dot in this figure is proportional to the connectivity of the respective habitat patch Stochasticity and space in population dynamics

Otso Ovaskainen Ace North senior researcher PhD -student

his year we extended our previous work developed earlier (e.g. Cornell and Ovaskainen on ecological dynamics to evolutionary 2008). Second, we took the approach of adaptive Tdynamics, focusing on how dispersal evolves dynamics by letting the mutation rate go to zero, under spatiotemporally varying environmental in which case the dynamics are characterized by a conditions. We considered a family of individual- rare mutant attempting to invade a monomorphic based models which all collapse to the same resident population. To analyze invasion rates for non-spatial mean-field limit of logistic population stochastic and spatial individual-based models, we growth. The models differ in terms of the process collaborated with Stephen Cornell in developing that is affected by density dependent competition perturbation theory for eigenvalues (Figure 1). (fecundity, establishment or mortality), and the process that is affected by habitat quality (fecundity, establishment, mortality, carrying capacity or none of these). We first assumed a finite mutation Collaborators rate for the genes affecting dispersal behavior, in which case the ESS (evolutionary stable strategy) is Dr. Stephen Cornell, Leeds University, UK. not a single genotype but a stationary frequency distribution of genotypes. The ESS frequency Key publications can be computed using the mathematical theory Cornell, S. J. and Ovaskainen, O. (2008). Exact Fig. 1. A comparison between simulations and asymptotic analysis for metapopulation dynamics theory on predicting (A) the invasion rate of a rare on correlated dynamic landscapes. Theoretical mutant attempting to invade a monomorphic resident Population Biology, 74, 209-225. population, and (B) the spatial pattern of invasion (mutant-resident correlation function in Fourier space). A B

22 23 Biodiversity informatics and conservation

Atte Moilanen Mar Cabeza Anni Arponen senior researcher senior researcher PhD -student

Heini Kujala Joona Lehtomäki PhD -student PhD -student

Undergraduate student: Johanna Eklund

n this group we develop concepts, theory, modeling, including modeling of species richness algorithms, methods and software for the needs and turnover of community composition, is a Iof spatial conservation prioritization, aiming at way forward in conservation planning at data- benefits for conservation due to an improved ability poor areas of the world. The idea is that species- to put ecologically justified plans and numbers rich and diverse areas can be identified without on the political planning table. The first part of explicit knowledge of which species occur and our work is to understand factors that explain where. Following conceptual understanding of this distributions of species and communities. We then approach in 2007, we have this year advanced with apply optimization, decision theory and uncertainty implementation and applications. analysis on top of state-of-the-art ecological modeling, with the aim of identifying efficient and Habitat loss is reducing conservation opportunities reliable conservation decisions. Major components globally. Another focus of our present work is of our work include species distribution, community dealing with dynamic landscapes and habitat loss and metapopulation modeling, spatial optimization in spatial reserve planning. This is important in the and methods for dealing with spatial connectivity, context of limited yearly budgets and sequential methods for dealing with habitat loss rates, climate- reserve selection, in which the reserve network change considerations, multi-action conservation is gradually built over several years. Habitat loss planning and applications to habitat restoration. We influences sequential planning, because sites may be work on applications together with collaborators on lost before they can be purchased, which should be all continents. accounted for. Another case of dynamic landscapes is species distribution shifts caused by climate Combination of community modeling and species- change. Dealing with climate change in conservation level modeling in reserve selection is one of our planning is particularly difficult due to multiple present main lines of investigation. Community highly uncertain processes that must be accounted

24 Fig. 1. Planning forest conservation in Finland using high-resolution GIS data and the Zonation prioritization software. This work is the first part of Joona Lehtomäki’s recently started PhD, and it has high practical relevance because it will be used to help target a 10,000 hectare expansion of present conservation areas of Metsähallitus (the Finnish Forest and Park Service). Panel A shows the conservation value of forests in Southern Finland with state-owned forests forced in as top priority. Based on the top fraction of valuable forests, panel B shows how the existing reserve network (in black) can be extended with new areas (in violet) of high quality and connectivity. for. Together, habitat loss and climate change are because different species benefit from different expected to cause large-scale shifts in species actions, and there may be tradeoffs between species. distributions as well as numerous extinctions, and The spatial multi-action planning problem with counteracting these processes is of highest priority habitat loss is already computationally a very hard in conservation planning. problem, - multimodal, nonlinear, stochastic, and high-dimensional. Dealing with landscape dynamics links to multi- action conservation planning, in which a site can With respect to the emphasis of recent and ongoing be treated in several alternative ways including work, we have (i) made big advances around the use different levels of protection, restoration or habitat of community modeling in conservation planning, maintenance. Multi-action planning allows a (ii) developed two methods for spatial conservation unified view of the landscape, with different actions planning accounting for climate change, (iii) out- (or lack of actions) influencing the occurrence lined a concept framework for conservation prioriti- of biodiversity features – multi-action planning zation, and (iv) participated in multiple high-profile allows for maximizing retention and persistence applications of the Zonation method. Additionally, of biodiversity across the entire landscape, not (v) we have contributed heavily to an edited volume only in conservation areas. One main variant of about spatial conservation prioritization, (vi) Zona- the multi-action planning problem is targeting of tion v2 has been released (see www.helsinki.fi/biosci- habitat restoration effort. This is particularly relevant ence/consplan), (vii) we have obtained funding from for example in central Europe, where there is little the SCALES EU project,(viii) we have arranged teach- land remaining that could be protected, but rather, ing of conservation biology at Madagascar, (ix) work conservation effort is increasingly directed towards on climate change issues has intensified, and (x) Mr. restoration of degraded habitats. The multi-action Joona Lehtomäki has started his PhD concerning a planning problem is computationally difficult, topic of high relevance in Finland (see figure).

25 Collaborators Key publications

National Arponen, A., Moilanen, A., and S. Ferrier. (2008). A successful community-level strategy for Finnish Environment Institute, Dr Risto Heikkinen, conservation prioritization. Journal of Applied Dr Juha Pöyry, Dr Mikko Kuussaari, and Dr Pekka Ecology, 45, 1436-1445. Punttila. Moilanen, A., K. A. Wilson, and H.P. Possingham, Finnish Forest Research Institute; Prof. Erkki Tomppo editors. (2009). Spatial Conservation Prioritization: and Dr Juha Siitonen Quantitative Methods and Computational Finnish Forest and Park Service: Dr Jussi Päivinen and Tools. Oxford University Press, in press, Panu Kuokkanen. scheduled for 5/2009 (see www.oup.com/uk/ Finnish Museum of Natural History; Prof Risto catalogue/?ci=9780199547777). Väisänen et al. Kremen, C., A. Cameron, A. Moilanen, S. Phillips, C. D. Thomas, et al. (2008). Aligning conservation International priorities across taxa in Madagascar, a biodiversity hotspot, with high-resolution planning tools. ACERA, Australian Centre of Excellence in Risk Science 320, 222-226. Analysis, directed by professor Mark Burgman; Dr Leathwick, J. R., Moilanen, A., Francis, M., Elith, J., Jane Elith, Dr Peter Vesk Taylor, P. Julian, K. and T. Hastie. (2008). Novel CERF/AEDA Commonwealth Research Hub in Applied methods for the design and evaluation of marine Environmental Decision Making, University of protected areas in offshore waters. Conservation Queensland, Australia, directed by professor Hugh Letters 1, 91-102. Possingham; Dr Kerrie Wilson, Dr Hedley Grantham Pressey, R.L., Cabeza, M., Watts, M.E., Cowling, R.M., Biodiversity and Global Change Laboratory, directed Wilson, K.A., (2007). Conservation planning in a by Prof Miguel Araújo, Natural History Museum, changing world. TREE 22, 583-592. Madrid Lab of Prof Chris D. Thomas, University of York; Dr Barb Anderson, Dr Jenny Hodgson et al. MACIS EU project (Minimization of and Adaptation to Climate Change Impacts on Biodiversity); Prof Miguel Araújo, Dr Wilfried Thuiller Prof John Leathwick, NIWA, New Zealand Prof Simon Ferrier, CSIRO, Australia Prof Bob Pressey, James Cook University, Australia Prof Claire Kremen, University of California, Berkeley Prof Ralph McNally, Dr Jim Thomson, Monash University, Australia Dr Ascelin Gordon, Dr Sarah Bekessy, RMIT, Royal Melbourne University of Technology Dr Simon Ferrier, Dept. Environment and Conservation, Australia. Dr John Leathwick, NIWA, New Zealand Dr Kerrie Wilson, Nature Conservancy, Australia

Fig. 2. SCIENCE HIGHLIGHT, cover of Science Cover of Zonation 320: Kremen, et al. (2008); worlds largest spatial v2.0 software manual, conservation prioritization study to date: expansion released in spring of protected areas of Madagascar using data for 2315 2008. endemic species modeled across Madagascar at a sub- 1 km2 resolution. Done using the Zonation software.

26 Brown bear project

Janne Sundell Jonna Katajisto Undergraduate students: post doc post doc Sanja Heikkilä, Piritta Korhonen

ur management oriented brown bear their location and concealment varies with project is based on long-term radio-tracking human influence. Otogether with novel GPS-GSM data. We • collection of insects associated in bear dung study factors affecting the distribution of bears and in order to study how the structure of insect their behaviour close to humans, with an objective community varies with the age and location to develop techniques and models that would of the dung, the diet of the bear, and between help in the management of our expanding bear seasons. populations. Current work includes • study on the performance of the GPS-GSM bear collar; special focus on alignment and • development of the habitat weighted method habitat for analysing home ranges and exploration of • collaboration with Scandinavian Brown Bear possibilities for using diffusion based approach Research Project in a study in which fresh for the same purpose. tracks of GPS marked bears are followed • the study on bear behaviour in contacts with with the help of dogs, and marks of different humans. More experiments in which the resting behaviours, such as foraging, bedding, and male bears have been approached on foot marking of territory, are recorded. are conducted to increase the reliability of our results. We have also altered the modes of approach, for example, by changing the Collaborators number of persons involved. • investigation of the characteristics of bears’ Jon Swenson, Sven Brunberg & Ole-Gunnar Støen, day-time resting beds to see, for example, how Scandinavian Brown Bear Research Project Ilpo Kojola, Finnish Game and Fisheries Research Institute Thorsten Wiegand, UFZ Centre for Environmental Research Leipzig

Key publications

Katajisto, J.K. and Moilanen, A. (submitted). Accounting for environmental heterogeneity in estimating animal home ranges: a habitat weighted kernel method. Katajisto, J., A. Moilanen, T. Wiegand, and J. E. Swenson. (submitted). Effects of targeted © Sanja Heikkilä harvesting on Scandinavian brown bears. Sundell, J., Kojola, I. And Hanski, I. (2006). A new GPS-GSM -basedmethod to study behaviour of Fig. 1. Above-ground winter den of the brown bear in brown bears. Wildlife Society Bulletin, 34, 446-450. South-Savo.

27 Biology of an old adaptive radiation: Evolutionary ecology of dung beetles in Madagascar

Ilkka Hanski Luisa Orsini Heidi Viljanen professor post doc PhD -student

Pierre Rahagalala Helena Wirta PhD -student PhD -student

Undergraduate student: Mirja Miinala

adagascar has a large fauna of ca 300 Second, a modeling study will be started combining endemic dung beetles, including two speciation-extinction dynamics and the dynamics Mmain radiations, the endemic subtribe of species’ geographical ranges, stimulated by our Helictopleurina with 65 species and several empirical results for Malagasy beetles. clades in the wide-spread tribe Canthonini with ca 170 species in Madagascar in several endemic Highlights for 2008: genera. This project was started in 2002. So far we have made good progress in reconstructing • Helena Wirta completed a molecular phylogeny phylogenies, in developing good understanding of the genus Nanos, documenting complex of the population ecology and breeding biology phylogenetic and phylogeographic patterns of individual species, and in accumulating a with cryptic species and introgression. comprehensive data base on the geographical ranges of the species. In 2009, two new lines • Pierre Rahagalala and others have documented of research will be initiated. We will sequence the structure of the dung beetle community the transcriptome of Nanos viettei (Canthonini), inhabiting cattle dung in open areas. This which belongs to an interesting group of about community is species-poor in comparison with 10 species. In the next stage a large number of comparable communities in mainland Africa, genes will be sequenced for most of the other apparently due to lack of native ungulates in species to study patterns in molecular variation, Madagascar and the relatively recent (1,500 yrs cryptic species, hybridization and introgression. ago) introduction of cattle to Madagascar.

28 1212 I. Hanski et al. Letter

(a) 5 (b) 5 (c) 5

4 4 4

3 3 3

2 2 2

1 1 1 Percentage sequence difference Percentage 0 0 0 024681012 024681012 024681012 Distance Distance Distance (d) (e) (f) 5 5 5

4 4 4

3 3 3

2 2 2

1 1 1 Percentage sequence difference Percentage 0 0 0 024681012 024681012 024681012 Distance Distance Distance

Figure 2 PercentageFig. 1. These sequence results difference demonstrate between that pairs the three of individuals species of Helictopleurus against the corresponding that have shifted geographical to use cattle distance (in degrees, correspondingdung to 112 in open km at habitats the equator). since Upperthe introduction row: Helictopleurus of cattle quadripunctatus to Madagascar(a), 1,500Helictopleurus yrs ago marsyas have greatly(b) and expandedHelictopleurus neoamplicollis (c). The opentheir symbols geographical in (b) are ranges for pairs following of individuals resource involving shift, apparentlyH. marsyas becauseand Helictopleurus resource shift nicollei relaxed(see text). resource Lower row: Helictopleurus unifasciatus (d),competition.Helictopleurus The perrieri vertical(e) and axisNanos gives clypeatus the percentage(f; from Wirtasequence 2008). difference (COI) between pairs of individuals against the corresponding geographical distance (in degrees, corresponding to 112 km at the equator). Species different patternsthat have of molecular shifted to use diversity cattle dung in CO1. (upperHelictopleu- row): Helictopleurusincluded quadripunctatus in its diet in (a), multiple H. marsyas populations (b) and across the rus neoamplicollisH. neoamplicollisand H. marsyas (c).have The open very symbols limited haplotypein panel b are fororiginal pairs of range.individuals Nonetheless, involving the sister geographical species H. distribution of diversity andmarsyas a single and haplotype H. nicollei is. presentForest species across that Madagas- have not shiftedhaplotypes to use cattle suggests dung (lower that row):H. quadripunctatusH. unifasciatus (d),has H. recently car. The otherperrieri few (e) haplotypes and Nanos differ clypeatus by only (f). one or two expanded its range relatively rapidly towards north, where nucleotide changes from the common one, and the rare just a single haplotype has been sampled in many localities. Collaborators haplotypes mostly occur in the same region in north-western One might think that range expansions following resource Madagascar. This pattern is most parsimoniously explained Olivieror host Montreuil, shift are the commonplace, Paris National and Museum they may be so, but by the species having shifted to cattle dung within a small putativeof Natural examples History, areFrance mostly anecdotal. For instance, region in north-western Madagascar, where the haplotype Braschler & Hill (2007) suggested that current range diversity is presently highest, followed by rapid range expansion in the butterfly Polygonia c-album in northern Europe Key publications expansion. is facilitated not only by climate warming but also by altered The pattern in H. quadripunctatus suggests a different Wirta,host plantH., Wirta preference H., Orsini from L. andHumulus Hanski lupulus I. to the more process,• a gradualThree endemic change of Helictopleurus diet across much species of the have original widespread(2008). An andold adaptive common radiationUlmusglabra of forestand Urtica dioica. range, whichmanaged may have to shift covered to cattle most dung of southern in open Mada- Somewhatdung beetles stronger in Madagascar. evidence hasMolecular been reported for the gascar, whereareas. the A species molecular currently study showsby Ilkka high Hanski haplotype and tephritidPhylogenetic fly Tephritis and Evolution conura, which 47, 1076-1089. has been inferred to have diversity. Thisothers hypothesis demonstrates is consistent that two with species the ecology shifted and Hanski,colonized I., Wirta,Cirsium H., oleraceum Nyman,from T. and populations using Cirsium current resourceto the use new of resourceH. quadripunctatus. within a smallUnlike area the in other heterophyllumRahagalala,(Diegisser P. (2008). Resourceet al. 2006). shifts Genetic diversity was two species,northwestern which are hardly Madagascar, ever sampled followed with otherby rapid types significantlyin Malagasy lower dung in flybeetles: populations contrasting on C. oleraceum, and there of resourcerange than cattle expansion dung, H. across quadripunctatus the entireis island. attracted in wasprocesses less spatial revealed genetic by structure dissimilar in thespatial populations sampled genetic patterns. Ecology Letters 11, 1208- large numbers to human excrement and also to carrion both on this host species than on C. heterophyllum, suggesting range • Heidi Viljanen has completed her study on the 1215. in open areas and in dry forests. Apparently, H. quadripunct- expansion following host shift to C. oleraceum. Definite cases biology of Nanos viettei, which has a primitive atus has abreeding broader diet biology than among the other the cattlespecies dung-using in the of host shift-induced range expansion involve parasitic fungi species, andlarge it is tribe probable Canthonini. that cattle dung was gradually that have shifted to domesticated plants (Zaffarano et al. 2008)

� 2008 Blackwell Publishing Ltd/CNRS 29 Ecology of small rodents and their predators

Olivier Gilg Janne Sundell researcher post doc

Undergraduate students: Christina Church, Katri Korpela, Paula Lampinen, Netta Lempiäinen, Sanna Mäkeläinen, Hanne Vihervaara

he project concentrates on the small rodent syndromes and personality, and phase dependence population dynamics using both theoretical and in behavioural and life history traits in voles. OG Tempirical approaches with a special focus on has been working on arctic predator-prey model predation effects. Experimental studies have been and studied its sensitivity and behaviour and, for conducted in different scales; in the laboratory, example, in different scenarios of climate change. outdoor enclosures, and in the field. Field studies He also used this model to compare and discuss are done in Finland and in Greenland. We have regional differences documented for the lemming studied, for example, large-scale spatial dynamics dynamics in Northeast Greenland. of voles and their predators and the numerical and functional responses of the predators of voles Collaborators and the collared lemmings. More recently JS has conducted research on the apparent competition Pekka Helle, Finnish Game and Fisheries Research and antipredatory behaviour of voles and their Institute consequences at the population and community Heikki Henttonen and Otso Huitu, Finnish Forest level. Undergraduate projects completed in 2008 Research Institute have focused on spatio-temporal aggregation, Benoit Sittler, University of Freiburg, Germany escape tactics of voles and weasel hunting Hannu Ylönen, University of Jyväskylä behaviour, and infanticide in voles. Ongoing projects include studies on the predation and Key publications seasonal effects on the reproduction, behavioural Fig. 1.The bank vole (Myodes glareolus) is an Trebaticka, L., Sundell, J., Tkadlec, E. & important species in boreal vole communities. Ylönen, H. (2008). Behaviour and resource use of two competing vole species with shared predation risk. – Oecologia, 4, 707-715. Eccard, J. A., Pusenius, J., Sundell, J., Halle, S. & Ylönen, H. (2008). Foraging patterns of voles under heterogeneous avian and uniform mustelid predation risk. - Oecologia, 4, 725-374. Sundell, J., Trebaticka, L., Oksanen, T., Ovaskainen, O., Haapakoski, M. & Ylönen, H. (2008). Predation on two vole species by a shared predator: antipredatory response and prey preference. - Population Ecology, 50, 257-266. Sundell, J. & Ylönen, H. (2008). Specialist predator in multi-species prey community: boreal voles and weasels. - © Janne Sundell Integrative Zoology, 3, 51-63

30 Butterfly expansion and climate change

Varpu Mitikka PhD -student

© Mia Valtonen emperate butterflies have been shown to Fig. 2. Map butterfly Araschnia( levana). respond to warming temperatures by expand- Ting their distribution ranges northwards. This project examines the influence of climate change regional pattern in allelic frequency of the metabolic and other associated factors on the expansion of enzyme PGI (phosphoglucose isomerase) (see the European map butterfly (Araschnia levana) in figure) which has in previous studies shown to be Finland. associated with butterfly flight metabolic rate and dispersal rate, flight capacity and dispersal rate. A climate-envelope model prediction of the species’ present distribution (for the period 2000-2004) The field observations from a mark-recapture failed to predict the regional distribution, but nearly experiment and land-cover data are used in a model correctly predicted the northern range limit. The to describe the habitat preference of the species. expansion occurs in two largely separated expansion The influence of habitat structure on the expansion fronts in southern and eastern Finland, with a will be further studied with model simulations. faster rate in the east. Regional differences in the expansion may be due to many factors. We found a Supervisors: Ilkka Hanski & Risto Heikkinen

Collaborators

Timothy Carter, Stefan Fronzek, Risto. K. Heikkinen and Juha Pöyry, Research Department, Finnish Environment Institute, Helsinki Miska Luoto, University of Oulu Kimmo Saarinen, South Karelia Allergy and Environment Institute, Tiuruniemi

Key publications

Mitikka, V., Heikkinen, R. K., Luoto, M., Araujo, M. B., Saarinen, K., Pöyry, J. & Fronzek, S. (2008). Predicting range expansion of the map butterfly in Northern Europe using bioclimatic models. Biodiversity and Conservation, 17, 623-641.

Fig. 1. Two main alleles a (in green) and b (in blue) were identified in PGI. The frequency of the a allele was higher in the eastern Finland than in the southern Finland and Estonia (F = 12.9, p = 0.001, Pearson X2 = 468), N = 234 individuals, 9, 5, and 4 populations were sampled in the above mentioned study areas

31 Spatially structured coevolution

Evolutionary epidemiology

• This year marks a milestone in the long-term study of the interaction between Po. plantaginis and Pl. lanceolata as we’ve begun the massive Anna-Liisa Laine parallel sequencing (454 sequencing) project for Po. plantaginis with the aim of developing post doc Single Nucleotide Polymorphism (SNP) markers. The developed markers enable a wide range of experimental approaches that require the Undergraduate students: genetic recognition of pathogen strains. They Riitta Ovaska, Terhi Lahtinen will also be used to study the population genetic structure of Po. plantaginis. The long-term research aim is a synthesis of the molecular and epidemiological data. Collaborator: Dr. Mikko Frilander (University of he aim of this work is to show how ecology Helsinki) drives the basic processes by which pathogens evolve, and to understand how evolutionary T • This year we completed a spatio-temporal and epidemiological dynamics combine to produce the patterns of pathogen occurrence we observe. model within a Bayesian framework of the The large natural metapopulation of the fungal regional dynamics of Po. plantaginis at the scale pathogen Podosphaera plantaginis infecting host of the Åland islands. The model allows us to es- plant Plantago lanceolata in the Åland Islands has timate key features of the epidemiology of Po. proved an ideal system this purpose. Previous and plantaginis - dispersal distances, areas of suc- ongoing research have demonstrated rapid ongoing cessful overwintering, and encounter intensities coevolution between these species despite the between the host and the pathogen (Fig. 1). high turnover rate of local pathogen populations Collaborators: Prof. Ilkka Hanski (University of Highlights for year 2008 were: Helsinki), Dr. Samuel Soubeyrand (INRA at Avi- gnon, France), Prof. Antti Penttinen (University of Jyväskylä)

Fig. 1. Smoothed posterior estimates of overwintering (left) of Po. plantaginis, and of its encounter rate (right) with Pl. lanceolata in Åland in 2001-2006.

32 Parasite evolution in heterogeneous environments

• We reviewed recent empirical and theoretical advances on how polymorphism is maintained in host-parasite interactions to conclude that spatially heterogeneous selection is sufficient to fulfil the conditions required of stable polymorphism. Collaborator: Dr. Aurélien Tellier (University of Münich, Germany)

• This year we went below ground to see how mycorrhizal symbionts may play into the interaction between Po. plantaginis and Pl. lanceolata. Preliminary results indicate that mycorrhizal colonization significantly reduces infection severity of the plants, and that plant genotypes differ significantly in their affinity to the mycorrhizal association. In the future we will further explore spatial variation in the mycorrhizal community associated with Plantago, and study how it may generate spatially divergent coevolutionary selection. Collaborator: Dr. Minna-Maarit Kytöviita (University of Jyväskylä)

• Experimental work on co-infection, i.e. when ©John N. Thompson the same host plant is infected by several different pathogen strains, demonstrated that the outcome is highly context dependent. The Fig. 2. Lithophragma cymbalaria. interaction among the strains appears to be mediated by the host, and the outcome ranged from an increase to a reduction in fitness of the subsequent infection compared to its performance as a single infection, depending on the environment. Future work aims to unravel the underlying mechanisms generating such variable outcomes of co-infection. Key publications

Coevolution at range margins Soubeyrand, S., Laine, A.-L., Hanski, I. & Penttinen, A. (In review). Spatio-temporal structure of host- • In a study of the widespread interaction pathogen interactions in a metapopulation. between moth Greya politella and its Laine, A.-L. & Tellier, A. (2008).Heterogeneous Lithophragma host plants in the northwest selection promotes polymorphism maintenance in US we are currently studying whether host-parasite interactions. Oikos 117, 1281-1288 coevolutionary selection varies at the different Laine, A.-L. (2008) Temperature-mediated patterns hierarchical levels when moving from individual of local adaptation in a natural plant-pathogen flowers of a plant to the plant level in metapopulation. Ecology Letters 11, 327-337 populations where the moth has shifted onto a Laine, A.-L., Thompson, J. N. & Thompson, J. F. (In novel host, L. cymbalaria (Fig. 2). review). Mutualistic and antagonistic pollinating Collaborator: Prof. John N. Thompson (University floral parasites affect seed set on a shared of California at Santa Cruz, US) hostplant.

33 Supporting personnel

y main responsibil- Mity is to maintain the computers and the www- y main task is to pages of the group, layout Mtake care of the publications and produce Glanville fritillary (Me- figures for different pur- litaea cinxia) laboratory poses. I also help with all populations and assist kind of practical things in researchers in their field the group. On maternity and laboratory experi- Camilla Ekblad leave from 10/2008. ments. I’m working year around in the new re- research secretary Suvi Ikonen search building at Lammi Biological Station. research assistant am taking care of data I recording during the Glanville fritillary survey and data extraction from facilitate the labo- the database for differ- I ratory work of the ent projects. I am also molecular ecology team programming user inter- by providing the neces- face and data handling sary materials, chemicals components for software and enzymes. Extracting, Evgeniy Meyke produced in the group. amplifying and sequenc- research assistant ing DNA is an essential part of my tasks as well as processing samples for y main duties screening of DNA micro concern the group’s M satellites. I also process budget and personnel Toshka Nyman samples for SNP (single administration. In addition laboratory technician nucleotide polymor- to that I am taking care of phism). various other things, for example organising meet- ings and trips. On maternity leave from 9/2008. joined MRG at the end Tuuli Ojala I of August 2008 to take research secretary care of goup’s budget and personnel related issues during Tuuli’s maternity have been organizing leave. I also take care of I and supervising the several minor tasks, like annual spring and autumn organising trips and visits.. surveys of Glanville fritillary butterfly in the Åland Marika Lilja islands. I also work with all computer related things in research secretary the group from maintenance to photoshopping of figures. Sami Ojanen project coordinator

34 Synopsis of the year 2008

Publications Theses Articles and book sections Books Articles, interviews and programs in public media

Honours, awards and memberships

Conferences and seminars

Work abroad

Visitors

Teaching

Budget

Prospects for the year 2008 Publications

MSc theses

Christina Church, MSc: Spatio-temporal aggregation in voles

ispersion describes the spatial distribution of I found that habitat selection of the bank vole and Dorganisms. We can recognize different main the field vole was mainly affected by the time of year types of dispersion. The type of dispersion playing and population density. There was a clear pattern a key role in this research is aggregated dispersion. indicating that both spcecies moved to inferior By investigating aggregated dispersion of voles and habitat types when the population density was high. shrews, we achieve valuable information on their In fields, which are the main habitat type for field basic biology. Voles and shrews are a crucial part voles, the density of the bank vole was never very of the northern ecosystem and a part of the diet of high. This suggests competition between these two many predators. The study of dispersion of small species. The results on the common shrew were not mammals has an important role in understanding as clear, but common shrew is a habitat generalist in the relationship between small mammals and their contrast to the vole species being studied here. predators. Especially for those predators specialiced in small mammals, the dispersion of voles can be of All species studied were aggregated in space great importance. when taking the species as a whole but there were differences between functional groups in the The purpose of this master’s thesis was to study population. For example the territorial female bank whether the bank vole, the field vole and the voles showed a segregated distribution where as common shrew are aggregated in space and time. immature individulas of the same species were The main focus of this study was on voles. I studied aggregated. The bank vole and the field vole were aggregation of these species on several hierarchical found to have a segregated distribution in relation levels. I also studied whether there were differences to each other, which again suggests competition in the distribution of the species in different habitats between these species. Aggregation in time or site and how population density affected their spatial tenancy was observed specially in the bank vole, distribution. I compared the two vole species and which lives in quite stabile habitats. Habitat stability the common shrew with each other and investigated is known to be a key factor influencing site tenancy. if there were differences in their dispersion and how these three species were distributed in relation to Based on my research, I can conclude that the each other. I also studied if for example the sex or main factors influencing the spatial aggregation the time of year affected the dispersion of these and aggregation in time in small mammals were: small mammals. species, sex and sexual state of an individual, habitat, population density and interactions I had a very large small mammal data, collected between species, competition being a representative using the small quadrat method, in my use. The example. trappings were performed during 1996-2000 on six islands which were located on three large lakes in the middle part of Finland, Lake Saimaa, Supervisors: Otso Ovaskainen & Janne Sundell Lake Päijänne and Lake Pielinen. I conducted the aggregation analyses with Toast 1.0 beta –program, with which it is possible to measure aggregation using relative propabilities. Before this study there has been only little research done on vole aggregation and also the studies concerning habitat selection have usually been based on much smaller data. All things considered, I believe that this study brings a new perspective to the study of small mammals.

36 Bess Hardwick, MSc: National Oak Gall Wasp Survey – mapping with parabiologists in Finland

parabiologist is an assistant in biological The survey produced data on 16 oak cynipid species, A research without formal academic training. In two of which were new to Finland. These new this study, I use volunteer parabiologists to conduct records bring the total number of species recorded a national survey of oak cynipid gall wasps, tribe in Finland up to 21. The centre of oak cynipid Cynipini (Hymenoptera: Cynipidae). My objectives species richness was in the southwest, but some were to produce an updated list of Finnish oak species were found as far north as Ostrobothnia. cynipid species, to map the distribution of these Within some species, different generations occurred species, and to examine factors that could have an on different individual trees. For this reason, I effect on local species richness and on the regional separated the generations in the distribution maps distribution and local abundance of individual and statistical analyses. Against a backdrop of large- species. I also wanted to examine whether the two scale clines in species richness, local species richness generations of a species tend to occur on the same was affected by the number of neighbouring oaks or on different individual trees. and by tree circumference. Large oaks that are surrounded by many other oaks support the most Empirical data on local gall wasp assemblages were species-rich communities of oak cynipids, and the collected by over a hundred volunteers from all local incidence of certain species and of generations over Finland, mostly people with a general interest within species exhibit similar patterns. in nature. Since most gall wasps occur as two discrete generations, volunteers sampled oak trees To conclude, oak cynipids are suitable targets for twice during the summer of 2007, with the spring sampling by parabiologists, and this survey has round encompassing 218 trees and the autumn improved our knowledge of the Finnish oak cynipid one 180. From each tree, five half-metre branches fauna. Oak cynipids have successfully spread beyond were examined, followed by a ten-minute scan of the host plant’s narrow natural distribution and the rest of the branches. The recovered samples colonised the planted trees in the north. Since large were then sent to me for identification. Based on oaks close to other oaks exhibit the most diverse tree-specific information supplied by the volunteers, gall wasp communities, the natural old oak forests in I used generalised linear models to examine the the southwest of Finland are particularly important effects of the number of neighbouring oaks, of to these species. oak circumference and of sampling location on the species richness of gall wasps, on the local incidence Supervisor: Tomas Roslin of species and on their local abundance.

Sanja Heikkilä, MSc: Performance of a GPS–GSM-method and its application in studying brown bear behaviour in human encounters

urrently in Finland, the brown bear (Ursus An additional advantage of the GPS is its great Carctos arctos) is returning to areas it has been accuracy in comparison with other methods. When absent for decades. Presence of the bear is of a combined with GSM (Global System for Mobile great concern to many people living in, or using, Communication) technology, as in this research, these areas. To form a basis on which brown bear it enables reception of the location data without mitigation, management and conservation methods a great time lag, and thus plotting positions on a can be built on, knowledge and understanding map as they arrive. This is of a great convenience about animals’ movements, habits, and behaviour when tracking the movements of the bear and the is of fundamental importance. Approaching approaching persons involved in the experiments. experiments on bears provide a way to gain fact- With any method, prior to its actual implementation, based information on brown bear reactions in it is essential to determine its functioning in human encounters. Recent, automatic, satellite- different environments and conditions. In earlier based tracking methods such as Argos or GPS studies, various factors have been proven to have (Global Positioning System) potentially cover an effect on the success and accuracy of the GPS- the entire planet and make collection of animal locations. In this study, I determine the effects location data feasible year-round, 24 hours a day, of tree density and canopy cover, position of the in all terrain, and without disturbing the animals. GPS-antenna, movement, and location interval on

37 the precision, accuracy, and success of location having an effect on the fix rates. The best interval attempts of four GPS–GSM bear collars and one for the approaching experiments, as a compromise GPS–GSM mobile phone. In addition to testing between various factors, was 30 seconds. For the conducted on stationary bear collars on open sites, margin of error to be taken into account in GPS the behaviour of both the moving and the resting locations, ± 6 metres was recommended. In case of bear was imitated by carrying the bear collars dense habitats, the margin should be wider. through different habitats, and by placing the collars In three experiments out of four the bear stayed in habitats bears seem to favour as their day-rest more or less immobile for the duration of the sites. The functioning of the bear collar in practice experiment. In one experiment the bear first was tested in experiments carried out to study bears remained immobile but after the approachers closed that were approached by humans. The aim was to in to a distance of 44 metres, the bear escaped. Due discover the optimal tracking solutions for the future to the small amount of experiments, conclusions experiments. about the bear’s behaviour can not be drawn. The GPS–GSM-method proved to be fairly reliable and The density and the canopy cover had the most accurate, and suited to its purpose for studying bear influence on the location error. The position of the behaviour. It enables the collection of more accurate antenna alone did not have a significant effect. The data on the reactions of the brown bear and also the total overall fix rate for GPS–GSM collars was 83 %, approachers, than has been possible so far. and for GPS–GSM phone 98 %. The GSM fix rates were higher, 99 % (collars) and 100 % (phone). The Supervisor: Janne Sundell GPS fix rate for the phone was 99 % and for collars 84 %. The location interval was the only factor

Eveliina Kallioniemi, MSc: Survival and growth of Glanville fritillary larvae: interaction between the effects of temperature and Pgi genotype

here is typically much variation in life-history non-f individuals. So far, the effects of Pgi genotype Ttraits among populations and genotypes within on larvae have not been studied. any one species. Such variation makes it possible for species to adapt to heterogeneous and changing In this study, I investigated the effects of the Pgi environments. Often there also exist trade-offs genotype on the survival and duration of the last between different traits or between different life- larval instar and pupal weight in the Glanville history stages. In the later case, a gene variant fritillary. Furthermore, I studied whether the effect that is beneficial at one life-history stage incurs a of Pgi genotype is dependent on population origin cost at another stage. One example of potentially (Åland in Finland, Estonia, and China), sex, and competing traits in insects is the duration of temperature (high, standard, and low). Treatment development and adult size. temperatures were selected to correspond to temperatures that larvae may encounter in the field. Temperature is one of the most important Additionally, larvae from Åland originated either environmental factors especially for invertebrates. from newly-established or old local populations, Ambient temperature affects growth and which are known to differ in numerous life-history development via its effects on metabolic rate. traits. Based on two SNPs (single nucleotide Variation in the metabolic enzyme phosphoglucose polymorphism), I divided the larvae to Pgi-non-f and isomerase (PGI) has been found to be strongly Pgi-f individuals. associated with variation in life-history traits. Different forms of PGI differ in their kinetic Larval development and survival was very different properties, and this can affect for instance flight among the temperature treatments: larvae did capacity, fecundity, and longevity. In the Glanville best in the high and worst in the low temperature fritillary butterfly (Melitaea cinxia), individuals with a treatment. Sex ratio among the adult butterflies was particular allele (Pgi-f) have been found to be able highly male biased. The experiment also revealed to fly under lower ambient temperatures and lay significant differences among the Pgi genotypes in larger egg clutches. Therefore, adult butterflies with both survival and growth. These differences were Pgi-f genotype are considered to be superior to Pgi- dependent on sex and population origin. The most

38 obvious difference was lower survival of the Pgi-f present result for larvae was the opposite since Pgi-f individuals in all treatments. individuals had lower survival in all treatments. This result may reflect a trade-off between the different Glanville fritillary females emerge on average a few life-history stages and could thereby contribute to days later and at larger size than males. It is hence the maintenance of genetic polymorphism in Pgi in possible that females are more sensitive to stressful this species. These results also underscore the need temperatures, which may at least partly explain the to study selection on a particular gene at every life- skewed sex ratio (due to disproportionate mortality history stage for comprehensive understanding of of females). The advantage that individuals gain the evolutionary forces acting on it. from developing fast or emerging at large size may vary among the sexes. Previous studies have shown Supervisor: Ilkka Hanski that Pgi-f individuals perform better as adults. The

Pirita Korhonen, MSc: The ecological and social carrying capacity of brown bears in Finnish game districts

he Ministry of Agriculture and Forestry has with stepwise linear regression how the attitudes Tset a goal that the brown bear (Ursus arctos) were related to the interest group, the geographic population in Finland should have a more uniform location and the recent trend in the bear population distribution. However, different areas are likely to size. Finally, I dissected the ecological and social have different habitat suitability for bears. Also the suitability of the GMDs in joint analysis. attitudes towards bears vary, which may lead to higher mortality rates in some areas e.g. because The GMDs of Lappi, Keski-Suomi, Kainuu and of poaching. As a consequence, bears may have Etelä-Savo have the largest areas with good quality surprisingly high mortality in high quality habitats habitat for bears, whereas the districts of Uusimaa, in areas where attitudes towards bears are negative. Ruotsinkielinen Pohjanmaa and Varsinais-Suomi This kind of areas may act as ecological traps. have very little suitable areas. The most common positive view points associated with bears were (i) In my master’s thesis I studied both the ecological the intrinsic value of bears, (ii) their value for nature and social suitability of Finnish game management and (iii) their value as game. The most common districts (GMD) for bears, which has not been done negative view points were (i) the fear of bears and before in joint analysis. I evaluated the ecological (ii) the damages to property. suitability by using PELCOM habitat classification and fitting to Finland a habitat suitability model According to my results especially the western parameterized with radiolocation data from Sweden. GMDs are both socially and ecologically poor I tested the predictive performance of the model for bears, which may hinder realising the goal in Finland by comparing with Spearman-rank of increasing the size of the bear population correlation the area-adjusted frequencies of bear in western Finland. In the southern Finland the observations within each habitat suitability class in attitudes towards bears are very positive, but there Pohjois-Karjala district to the corresponding rank is not much good quality habitat. The GMDs of of that class. The test indicated relatively good Pohjois-Savo, Etelä-Savo and Keski-Suomi function performance. I then defined the ecological suitability as bridge from east to west and south, and thus of the GMD as the total area of good quality habitat play important role in achieving more uniform within it. The social suitability I evaluated with help bear population in Finland. Indeed Etelä-Savo has of a questionnaire. Different interest groups were good suitability for bears but the GMDs of Pohjois- asked to list maximum three negative and three Savo and Keski-Suomi may act as ecological traps, positive points associated with bears and to give because they have good ecological suitability but those points a weight [-5, 5] according to their poor social suitability for bears. In Pohjois-Savo importance. I categorized the interest groups and there are also long water bodies and wide areas of the view points in meaningful groups and used human settlement, which may prevent spreading of the mean weight of the view points in each GMD the bear population. as definition for its social suitability. I also tested

39 Especially in those GMDs, where the fear of bears as important as the fear of bears. In the light of my was considered the most important point associated results the entire goal of more uniformly distributed with bears, e.g. Keski-Suomi and Pohjois-Savo, the bear population over Finland can be questioned: social suitability could be improved by investing both ecological and social suitability for bears seems to right kind of informing of the public. In those poor in western Finland and there is not much good districts, where the damages to property was the habitat for bears in southern Finland. Moreover, most important point, e.g. Lappi, the management some dispersal areas between these and the eastern actions should focus on preventing damages. areas with high number of bears may function as Investing to the prevention could improve the social ecological traps. acceptance of bears also in the GMDs of Keski- Suomi, Kymi, Etelä-Häme and Satakunta, where Supervisor: Jonna Katajisto the damages to property was considered almost

Anniina Mattila, MSc: Association Between Molecular Variation in the Phosphoglucose Isomerase (Pgi) Locus and Migration Propensity in the Glanville Fritillary Butterfly

abitat fragmentation produces patches of level examinations. Each butterfly and larva was Hsuitable habitat surrounded by unfavourable genotyped at the Pgi locus. The MRR data was matrix habitat. A species may persist in such a parameterised with two statistical models of fragmented landscape in an equilibrium between migration: the Virtual Migration Model (VM) and the extinctions and recolonizations of local the spatially explicit diffusion model. VM model populations, thus forming a metapopulation. predicted and observed numbers of emigrants Migration between local populations is necessary from populations with high and low frequencies for the long-term persistence of a metapopulation. of Pgi-f were compared. Posterior predictive data sets were simulated based on the parameters of the The Glanville fritillary butterfly (Melitaea cinxia) diffusion model. Lack-of-fit of observed values to forms a metapopulation in the Åland islands the model predicted values of several descriptors in Finland. There is migration between the of movements were detected, and the effect of populations, the extent of which is affected by Pgi genotype on the deviations was assessed by several environmental factors and variation in the randomizations including the genotype information. phenotype of individual butterflies. Different allelic forms of the glycolytic enzyme phosphoglucose This study revealed a difference in the effect of Pgi isomerase (Pgi) has been identified as a possible genotype on migration propensity between the genetic factor influencing flight performance and two sexes in the Glanville fritillary. The females with migration rate in this species. The frequency of a and males without the Pgi-f allele moved more certain Pgi allele, Pgi-f, follows the same pattern between habitat patches, which is probably related in relation to population age and connectivity as to differences in the function of flight in the two migration propensity. Furthermore, variation in flight sexes. Females may use their high flight capacity to metabolic performance, which is likely to affect migrate between habitat patches to find suitable migration propensity, has been linked to genetic oviposition sites, whereas males may use it to variation in Pgi or a closely linked locus. The aim acquire mates by keeping a territory and fighting of this study was to investigate the association off other intruding males, possibly causing them between Pgi genotype and the migration propensity to emigrate. The results were consistent across in the Glanville fritillary both at the individual and different movement descriptors and at the individual population levels using a statistical modelling and population levels. The effect of Pgi is likely to approach. be dependent on the structure of the landscape and the prevailing environmental conditions. A mark-release-recapture (MRR) study was conducted in a habitat patch network of M. cinxia Supervisor: Ilkka Hanski in Åland to collect data on the movements of individual butterflies. Larval samples from the study area were also collected for population

40 Sanna Mäkeläinen, MSc: Escape tactics of voles and predator’s response

here is an ongoing evolutionary arms race Escape reactions of vole species differed markedly Tbetween predators and their prey. Predation risk under weasel presence. Almost all field voles and unsuccessful predation develops prey’s escape escaped through the tunnel, but about 30% of tactics, but also predators become more effective in bank voles escaped by climbing. Field voles started evolutionary time scale. Small mustelids are adapted escaping even earlier than bank voles. Climbing to hunt rodents in their underground tunnels. The by voles made successful following of prey more least weasel (Mustela nivalis nivalis) causes most difficult for weasels. mortality in cyclic vole populations in Fennoscandia and voles have developed different antipredatory My results showed that climbing is an effective es- behaviours to reduce the probability of successful cape reaction, even if weasels can climb when they predation by the weasel. Several escape tactics have are motivated. Climbing can be beneficial because been documented in bank voles (Myodes glareolus), olfactory cues of climbing vole might be lost or but climbing in trees is assumed to be specialised because vole can attain more distance by jump- reaction for predation risk caused by the weasel. Re- ing down from the tree if also weasel is climbing. duced activity or escaping to underground tunnels is Because bank voles climbed more often than field more typical for field voles (Microtus agrestis), whose voles in my experiment and because weasels had escaping tactics are not so developed. difficulties to follow climbing voles it is likely that bank voles will survive better also in natural condi- The aim was to study experimentally escape tions under predation of their most deadly predator, tactics of bank voles and field voles and predator’s the least weasel, if there are possibilities for arboreal response to these tactics. Voles were tested in escape. Although field voles are easier to follow, laboratory in a tube system under a simulated they can pose a threat for small weasels because predation risk by a live least weasel. Vole had a of their larger size and more aggressive behaviour, possibility to escape the weasel either by climbing which has to be taken into account when consider- or trough a horizontal tunnel. After vole’s observed ing prey choice of weasels and success of the attacks escape reaction it was removed and the weasel was on the prey. released into the tube system in order to see if it could follow the vole. Supervisor: Janne Sundell

Markku Ojanen, MSc: Multitrophic interaction between plant, pathogen, herbivore and parasitoid

studied how infection by a plant pathogen on a H-groups used uninfected plant as food and HC- Ishared host plant may affect the parasitoid of an group were parasitized. Also, there were two treat- insect herbivore. The plant pathogen was hypoth- ment groups parasitized FCD- and HCD-groups in esized to cause changes in the quality of the host which the larvae were dissected in August in order to plant that may affect its herbivore and in turn have determine whether they were paratisized before dia- an impact on its parasitoid. The experiment was car- pause. Then total number of larvae in my experiment ried out with Powdery mildew Podosphaera plantag- was 540. The numbers of female wasps, which have inis, the host plant Narrow-leaf plantain (Plantago been used to parasitize butterfly larvae were 37 and lanceolata), larvae of an insect herbivore Glanville they came from 16 families. I searched for differences Fritillary Butterfly (Melitaea cinxia) and the parasitic between treatments by using following variables: wasp Cotesia melitaerum. I performed part of the development time of butterfly larvae from instar to experiment before diapause from June to August instar, number of butterfly larvae before and after 2005 and the part after diapause from April to May diapause, weight of butterfly larvae, number of wasp 2006 in Åland Islands in Finland. larvae inside host larva before diapause, number of adult wasps, weight of adult wasps and sex of adult I had nine families of butterfly larvae and I divided wasp. Statistical tests were done by using GLMM and each of them to six treatment groups. FC-groups GLM test. There was a statistically significant dif- used mildew-infected plant as food and they were ferences in the sex-ratio, which was female-biased parasitized by mated female wasps. F-groups ate in FC-groups and male-biased in HC-groups. This also infected plant, but were unparasitized. HC- and means that pathogen infection in herbivore food plant

41 increases number of female offspring or decreases References: male offspring. Laine, A-L. 2004. A powdery mildew infection on However, two things must be noticed. First, my a shared host plant affects the dynamics of the experiment was a pure laboratory experiment and Glanville fritillary butterfly populations. Oikos natural effects have been eliminated. My purpose 329-337. was to do also a field experiment, but it failed and thus I can not conclude on the effects of natural Supervisors: Saskya van Nouhuys and Anna-Liisa features in this study. Second, an earlier study (Laine Laine 2004) showed, that mildew-infection has negative effects for survival and development of butterfly larvae. I did not get statistically significant differences on the effect of infection on the insect herbivore.

Hanne Vihervaara, MSc: The effects of male infanticide and habitat fragmentation on the breeding success of the bank vole

nfanticide, the killing of conspecific young, oc- large habitat patch with male infanticide. There are Icurs in over 100 species of mammals, and it is two ways to achieve this; males, including infanti- commonly recognized as an adaptive behavioural cidal ones, avoid moving over the inter-patch matrix strategy which enhances the fitness of the perpetra- due to possible avian predation, thus nests in large tor. Infanticide is inhibited in several male rodent patches are more vulnerable to infanticide, or females species at the time when genetically related offspring in small patches do not move because of risk in the would be born, which is adaptive since males do not matrix area, therefore they stay closer to nest site and recognize their own offspring. It has been suggested protect the litter. that copulation alone is enough to inhibit infanticidal behaviour in male rodents. Infanticidal behaviour In the laboratotory experiment, I studied whether occurs in more than one third of bank voles (Myodes infanticidal male bank voles after successful mat- glareolus), which might have great effect on breeding stop the killing of pups. Infanticidal males were ing success and population dynamics of bank voles. paired with a female, and after the young were born Presumably infanticide is also inhibited in male bank the males’ infanticidal behaviour was studied several voles when their own young are born, although it is times. The hypothesis stated that paired infanticidal not studied in bank voles. males are inhibited from committing infanticide when presumed offspring are born. Also, it was assumed Habitat fragmentation degrades continuous natural that mating alone is sufficient for inhibition. habitats into smaller patches and affects usually negatively population dynamics in small mammals. The results of the enclosure experiment revealed that Habitat fragmentation also affects the spacing pat- the number of young was higher in the enclosures of terns of small mammals. Animals have to enlarge small patches, where females stayed closer to their their home ranges by crossing inter-patch area to get nests avoiding the matrix area, compared to the access to resources, or animals accept to stay in one enclosures of one large patch. The effect of infantici- patch, which reduces their movements. dal males on nest survival remained unclear probably because of the low survival rate of males in the enclo- I studied the combined effect of male infanticide sures. However, the results suggest that small-scale and experimental small-scale habitat fragmentation fragmentation might not affect necessarily deleterious on breeding success in bank voles in large outdoor on the population dynamics of the bank vole. enclosures (0.25 ha). The habitat of enclosures was manipulated by forming either one large or four In the laboratory experiment, 58 % of the infanticidal small habitat patches, which were surrounded by males halted the killing the young after successful a mowed inhospitable matrix area. Females with mating. Probably mating alone is not sufficient to a litter and either infanticidal or non-infanticidal inhibit the infanticidal behaviour completely. Pos- males occupied each enclosure during the experi- sibly besides copulation, also past association with ment. The movements of voles were observed with the mother is needed to the higher inhibition rate radio-tracking, and after the experiment all survived of infanticide, as it is found in some other species of young were counted. I expected that the number of rodents. survived young is higher in smaller, isolated patches without male infanticide and lowest in the single, Supervisor: Janne Sundell

42 Articles and book sections Evoluutio Nyt! (eds P. Portin & T. Vuorisalo). Kirja- Aurora, Turku. Hanski, I. (2008) Insect conservation in boreal forests. Arellano, L., León-Cortés, J.L., & Ovaskainen, O. (2008) Journal of Insect Conservation, 12, 451-454. Patterns of abundance and movement in relation Hanski, I. (2008). Metapopulation models. In Popula- to landscape structure: a study of a common scarab tion dynamics. Vol 3 of Encyclopedia of Ecology (eds (Canthon cyanellus cyanellus) in Southern Mexico. S.E. Jørgensen & B.D. Fath), pp. 2318-2325. Elsevier, Landscape Ecology, 23, 69-78. Oxford. Arponen, A., Moilanen, A., & Ferrier, S. (2008) A Hanski, I. (2008) Spatial patterns of coexistence of com- successful community-level strategy for conserva- peting species in patchy habitats. Theoretical Ecology, tion prioritization. Journal of Applied Ecology, 45, 1, 29-43. 1436-1445. Hanski, I. (2008) The world that became ruined - Our Cabeza, M., Arponen, A., & VanTeeffelen, A. (2008) cognitive incapacity to perceive large-scale and Top predators: Hot or not? A call for systematic long-term changes is a major obstacle to rational assessment of biodiversity surrogates. Journal of environmental policies. EMBO Reports, 9, S34-S36. Applied Ecology, 45, 976-980. Hanski, I. (in press). Incorporating the spatial configu- Cornell, S. & Ovaskainen, O. (2008) Exact asymptotic ration of the habitat into ecology and evolutionary analysis for metapopulation dynamics on corre- biology. In Spatial Ecology (ed S. Cantrell). CRC Press, lated dynamic landscapes. Theoretical Population London. Biology, 74, 209-255. Hanski, I. (in press). Metapopulations and spatial popu- Eccard, J.A., Pusenius, J., Sundell, J., Halle, S., & Ylönen, lation processes. In The Princeton Guide to Ecology. H. (2008) Foraging patterns of voles at heteroge- Princeton University Press, Princeton. neous avian and uniform mustelid predation risk. Hanski, I. (in press). The theories of island biogeogra- Oecologia, 157, 725-734. phy and metapopulation dynamics: Science marches Fischer, H.M., Wheat, C.W., Heckel, D.G., & Vogel, H. forward, but the legacy of good ideas lasts for a long (2008) Evolutionary origins of a novel host plant time. In Island Biogeography 40 years (eds R. Ricklefs detoxification gene in butterflies. Molecular Biology & J. Losos). Harvard University Press, Harvard. and Evolution, 25, 809-820. Hanski, I. (in press). Täpläverkkoperhonen populaatio- Franco, A.M.A., Anderson, B., Roy, D.B., Gillings, S., Fox, biologian mallilajina. In Sphinx Yearbook 2007-08, pp. R., Moilanen, A., & Thomas, C.D. (in press) Surroga- 45-54. cy and persistence in reserve selection: landscape Hanski, I., Wirta, H., Nyman, T., & Rahagalala, P. (2008) prioritisation for multiple taxa in Britain. Journal of Resource shifts in Malagasy dung beetles: contrast- Applied Ecology. ing processes revealed by dissimilar spatial genetic Frilander, M.J. (2008) Täpläverkkoperhonen liiti geno- patterns. Ecology Letters, 11, 1208-1212. miaikaan. Duodecim, 124, 2242-2243. Hodgson, J., Moilanen, A., & Thomas, C.D. (in press) Grantham, H.S., Moilanen, A., Wilson, K.A., Pressey, The responses of butterfly metapopulations to patch R.L., Rebelo, T.G., & Possingham, H.P. (2009) Dimin- connectivity and average patch quality are masked ishing returns for biodiversity data in conservation by successional habitat dynamics. Ecology. planning. Conservation Letters, 1, 190-198. Hottola, J. & Siitonen, J. (2008) Significance of wood- Gripenberg, S., Ovaskainen, O., Morrien, E., & Roslin, land key habitats for polypore diversity and red- T. (2008) Spatial population structure of a specialist listed species in boreal forests. Biodiversity and leaf-mining moth. Journal of Animal Ecology, 77, Conservation, 17, 2559-2577. 757-767. Koivisto, E., Huitu, O., Sundell, J., & Korpimäki, E. (2008) Gripenberg, S. & Roslin, T. (2008) Neither the devil Species-specific limitation of vole population growth nor the deep blue sea: larval mortality factors fail to by least weasel predation: facilitation of coexistence? explain the abundance and distribution of Tischeria Oikos, 117, 6-12. ekebladella. Ecological Entomology, 33, 346-356. Kremen, C., Cameron, A., Moilanen, A., Phillips, S.J., Haag-Liautard, C., Pederson, J., Ovaskainen, O., & Thomas, C.D., Beentje, H., Dransfield, J., Fisher, B.L., Keller, L. (2008) Breeding system and reproduc- Glaw, F., Good, T.C., Harper, G.J., Hijmans, R.J., Lees, tive skew in a highly polygynous ant population. D.C., Louis, E., Nussbaum, R.A., Raxworthy, C.J., Razaf- Insectes Sociaux, 55, 347-354. impahanana, A., Schatz, G.E., Vences, M., Vieites, D.R., Halme, P., Kotiaho, J., Ylisirniö, A.-L., Hottola, J., Jun- Wright, P.C., & Zjhra, M.L. (2008) Aligning conserva- ninen, J., Kouki, J., Lindgren, M., Mönkkönen, M., tion priorities across taxa in Madagascar with high- Penttilä, R., Renvall, P., Siitonen, J., & Similä, M. resolution planning tools. Science, 320, 222-226. (2008) Perennial polypores as indicators of annual Kremen, C., Cameron, A., Razafimpahanana, A., Moi- and red-listed polypores. Ecological Indicators, 9, lanen, A., Thomas, C.D., Beentje, H., Dransfield, J., 256-266. Fisher, B.L., Glaw, F., Good, T.C., Harper, G.J., Hijmans, Hanski, I. (2008). Evoluutio metapopulaatioissa. In

43 R.J., Lees, D.C., Louis, E., Nussbaum, R.A., Phillips, S.J., Moilanen, A., Possingham, H.P., & Wilson, K. (in press) Raxworthy, C.J., Schatz, G.E., Vences, M., Vieites, D.R., Spatial conservation prioritization: past, present and Wright, P.C., & Zjhra, M.L. (2008) Conservation with future. In Spatial Conservation Prioritization (eds A. caveats - Response. Science, 321, 341-342. Moilanen, K.A. Wilson & H.P. Possingham). Oxford Laine, A.-L. (2008) Temperature-mediated patterns of University Press, Oxford. local adaptation in a natural plant-pathogen meta- Moilanen, A., van Teeffelen, A., Ben-Haim, Y., & Ferrier., population. Ecology Letters, 11, 327-337. S. (2008) How much compensation is enough? Laine, A.-L. & Tellier, A. (2008) Heterogeneous selec- Explicit incorporation of uncertainty and time dis- tion promotes polymorphism maintenance in host- counting when calculating offset ratios for impacted parasite interactions. Oikos, 117, 1281-1288. habitat. Restoration Ecology, doi: 10.1111/j.1526- Leathwick, J.R., Moilanen, A., Francis, M., Elith, J., Taylor, 100X.2008.00381.x. P., Julian, K., & Hastie, T. (2008) Novel methods for Nicholson, E. & Ovaskainen, O. (in press) Conserva- the design and evaluation of marine protected areas tion prioritization using metapopulation models. In in offshore waters. Conservation Letters, 1, 91-102. Spatial conservation prioritization (eds A. Moilanen, Lewinsohn, T.M. & Roslin, T. (2008) Four ways towards H.P. Possingham & K.A. Wilson). Oxford University tropical herbivore megadiversity. Ecology Letters, 11, Press, Oxford, UK. 398-416. Niitepõld, K., Smith, A.D., Osborne, J.L., Reynolds, D.R., Marden, J.H., Fescemyer, H.W., Saastamoinen, M., Mac- Carreck, N.L., Martin, A.P., Marden, J.H., Ovaskainen, Farland, S.P., Vera, J.C., Frilander, M.J., & Hanski, I. O., & Hanski, I. (in press) Flight metabolic rate and (2008) Weight and nutrition affect pre-mRNA splic- Pgi genotype influence butterfly dispersal rate in ing of a muscle gene associated with performance, the field. Ecology. energetics and life history. Journal of Experimental O’Hara, R.B., Cano Arias, J.M., Ovaskainen, O., Teplitsky, Biology, 211, 3653-3660. C., & Alho, J. (2008) Bayesian approaches in evolu- Mitikka, V., Heikkinen, R. K., Luoto, M., Araujo, M. B., tionary quantitative genetics. Journal of Evolutionary Saarinen, K., Pöyry, J. & Fronzek, S. (2008) Predicting Biology, 21, 949-957. range expansion of the map butterfly in Northern Orsini, L., Corander, J., Alasentie, A., & Hanski, I. (2008) Europe using bioclimatic models. Biodiversity and Genetic spatial structure in a butterfly metapopula- Conservation, 17:623-641. tion correlates better with past than present demo- Moilanen, A. (2008) Generalized Complementarity and graphic structure. Molecular Ecology, 17, 2629-2642. Mapping of the Concepts of Systematic Conserva- Orsini, L., Wheat, C.W., Haag, C.R., Kvist, J., Frilander, tion Planning. Conservation Biology, early online, M.J., & Hanski, I. (in press) Fitness differences as- DOI: 10.1111/j.1523-1739.2008.01043.x. sociated with Pgi SNP genotypes in the Glanville Moilanen, A. (2008) Two paths to a suboptimal solu- fritillary butterfly (Melitaea cinxia). Journal of Evolu- tion - once more about optimality in reserve selec- tionary Biology. tion. Biological Conservation, 141, 1919-1923. Ovaskainen, O. (2008) Analytical and numerical tools Moilanen, A., Arponen, A., Stokland, J., & Cabeza., M. for diffusion based movement models. Theoretical (in press) Assessing replacement cost of conserva- Population Biology, 73, 198-211. tion areas: how does habitat loss influence priori- Ovaskainen, O., Cano Arias, J.M., & Merilä, J. (2008) A ties? Biological Conservation. Bayesian framework for comparative quantitative Moilanen, A. & Ball, I. (in press) Heuristic and approxi- genetics. Proceedings of the Royal Society B: Biologi- mate optimization methods for spatial conservation cal Sciences, 275, 669-678. prioritization. In Spatial Conservation Prioritization Ovaskainen, O., Luoto, M., Ikonen, I., Rekola, H., Meyke, (eds A. Moilanen, K.A. Wilson & H.P. Possingham). E., & Kuussaari, M. (2008) An empirical test of a Oxford University Press, Oxford. diffusion model: predicting clouded apollo move- Moilanen, A., Kujala, H., & Leathwick, J. (in press) The ments in a novel environment. American naturalist, Zonation framework and software for conservation 171, 610-619. prioritization. In Spatial Conservation Prioritization Ovaskainen, O., Rekola, H., Meyke, E., & Arjas, E. (2008) (eds A. Moilanen, K.A. Wilson & H.P. Possingham). Bayesian methods for analyzing movements in het- Oxford University Press, Oxford. erogeneous landscapes from mark-recapture data. Moilanen, A., Leathwick, J., & Elith, J. (2008) A method Ecology, 89, 542-554. for spatial freshwater conservation prioritization. Ovaskainen, O., Smith, A.D., Osborne, J.L., Reynolds, Freshwater Biology, 53, 577-592. D.R., Carreck, N.L., Martin, A.P., Niitepõld, K., & Moilanen, A., Possingham, H.P., & Polasky, S. (in Hanski, I. (2008) Tracking butterfly movements with press) A mathematical classification of conserva- harmonic radar reveals an effect of population age tion prioritisation problems. In Spatial Conservation on movement frequency. PNAS, 105, 19090-19095. Prioritization (eds A. Moilanen, K.A. Wilson & H.P. Paatero A.O., T.H., Happonen L.J., Olsson C., Palomäki Possingham). Oxford University Press, Oxford. T., Pajunen M.I., Meng X., Otonkoski T., Tuuri T.,

44 Berry C., Malani N., Frilander M.J., Bushman F.D. & tory response and prey preference. Population Savilahti H. (in press) Bacteriophage Mu integration Ecology, 50, 257-266. in yeast and mammalian genomes. Nucleic Acid Sundell, J. & Ylönen, H. (2008) Specialist predator in Research. multi-species prey community: boreal voles and Patterson, T.A., Thomas, L., Wilcox, C., Ovaskainen, O., weasels. Integrative Zoology, 3, 51-63. & Matthiopoulos, J. (2008) State-space models of Thomson, J.R., Moilanen, A., McNally, R., & Vesk, P. (in individual animal movement. Trends in Ecology and press) A quantitative method for prioritizing land- Evolution, 23, 87-94. scape revegetation. Ecological Applications. Pessa, H.K.J., Will, C.L., Meng, X., Schneider, C., Watkins, Trebaticka, L., Sundell, J., Tkadlec, E., & Ylönen, H. N.J., Perälä, N., Nymark, M., Turunen, J.J., Lührmann, (2008) Behaviour and resource use of two com- R., & Frilander, M.J. (2008) Minor spliceosome competing vole species under shared predation risk. ponents are predominantly localized in the nucleus. Oecologia, 157, 707-715. Proc. Natl. Acad. Sci., 105, 8655-8660. Turunen, J.J., Will, C.L., Grote, M., Lührmann, R., & Possingham, H.P., Wilson, K.A., & Moilanen, A. (in Frilander, M.J. (2008) The U11-48K protein contacts press) Accounting for habitat dynamics in conserva- the 5’ splice site of U12-type introns and the U11- tion planning. In Spatial Conservation Prioritization 59K protein. Molecular and Cellular Biology, 28, (eds A. Moilanen, K.A. Wilson & H.P. Possingham). 3548-3560. Oxford University Press, Oxford. Wahlberg, N. & Wheat, C.W. (2008) Genomic outposts Rahagalala, P., Viljanen, H., Hottola, J., & Hanski, I. (in serve the phylogenomic pioneers: Designing novel press) Assemblages of dung beetles using intro- nuclear markers for genomic DNA extractions of duced cattle dung in Madagascar. African Entomol- lepidoptera. Systematic Biology, 57, 231-242. ogy. van Nouhuys, S. & Kaartinen, R. (2008) A parasitoid Rayfield, B., Moilanen, A., & Fortin., M.-J. (in press) In- wasp uses landmarks while monitoring potential corporating consumer-resource spatial interactions resources. Proceedings of the Royal Society B, 275, in reserve design. Ecological Modelling. 377-385. Reudler Talsma, J., Biere, A., Harvey, J.A., & van van Nouhuys, S. & Laine, A.-L. (2008) Population Nouhuys, S. (2008) Oviposition cues for a special- dynamics and sex ratio of a parasitoid altered by ist butterfly: plant chemistry and size. Journal of fungal infected diet of host butterfly. Proceedings of Chemical Ecology, 34, 2102-1212. the Royal Society B, 275, 787-795. Reudler Talsma, J., Torri, K., & van Nouhuys, S. (2008) van Teeffelen, A. & Moilanen, A. (2008) Where and Host plant use by the Heath fritillary butterfly, Me- how to manage: Optimal allocation of alternative litaea athalia: plant habitat, species and chemistry. conservation management actions. Biodiversity Arthropod-Plant Interactions, 2, 63-75. Informatics, 5, 1-13. Rosenlew, H. & Roslin, T. (2008) Habitat fragmentation Vera, J.C., Wheat, C.W., Fescemyer, H.W., Frilander, and the functional efficiency of temperate dung M.J., Crawford, D.L., Hanski, I., & Marden, J.H. (2008) beetles. Oikos, 117, 1659-1666. Rapid transcriptome characterization for a nonmod- Roslin, T. & Salminen, J.P. (2008) Specialization pays el organism using 454 pyrosequencing. Molecular off: contrasting effects of two types of tannins on Ecology, 17, 1636-1647. oak specialist and generalist moth species. Oikos, Wheat, C.W. (in press) Rapidly developing functional 117, 1560-1568. genomics in ecological model systems via 454 tran- Roslin, T., Syrjala, H., Roland, J., Harrison, P.J., Fownes, scriptome sequencing. Genetica. S., & Matter, S.F. (2008) Caterpillars on the run - in- Wheat, C.W. & Watt, W.B. (2008) A mitochondrial- duced defences create spatial patterns in host plant DNA-based phylogeny for some evolutionary- damage. Ecography, 31, 335-347. genetic model species of Colias butterflies (Lepi- Saastamoinen, M. & Hanski, I. (2008) Genotypic and doptera, Pieridae). Molecular Phylogenetics and environmental effects on flight activity and oviposi- Evolution, 47, 893-902. tion in the Glanville fritillary butterfly. American Wirta, H. (in press) Complex phylogeographic pat- naturalist, 171, E701-E712. terns, introgression and cryptic species in a lineage Siitonen, J., Hottola, J., & Immonen, A. (in press) Dif- of Malagasy dung beetles (Coleoptera: Scarabaei- ferences in stand characteristics between brook- dae). Biological Journal of the Linnean Society. side key habitats and managed forests in southern Wirta, H. & Montreuil, O. (2008) Evolution of the Can- Finland. Silva Fennica. thonini Longitarsi (Scarabaeidae) in Madagascar. Soubeyrand, S., Laine, A.-L., Hanski, I., & Penttinen, A. Zoologica Scripta, 37, 651-663. (in press) Spatio-temporal structure of host-patho- Wirta, H., Orsini, L., & Hanski, I. (2008) An old adap- gen interactions in a metapopulation. tive radiation of forest dung beetles in Madagascar. Sundell, J., Trebaticka, L., Oksanen, T., Ovaskainen, O., Molecular Phylogenetics and Evolution, 47, 1076- Haapakoski, M., & Ylönen, H. (2008) Predation on 1089. two vole species by a shared predator: antipreda-

45 Books Hurme), 1/2008. Roslin, T. Rovaniemellä kerätään lantakuoriaisia. In Uusi Rovaniemi, 25.6. Moilanen, A. & Kujala, H. Roslin, T. Shovelling for beetles. In Helsinki University (2008) Zonation spatial Bulletin, (ed. Virve Pohjanpalo), 2/2008. conservation planning Roslin, T. Ötökkäsäilykkeitä Mynämäeltä. In Yliopisto, framework and software v. (ed. Virve Pohjanpalo), 9/2008. 2.0, User manual, Helsinki. Sundell, J. Etelä-Savossa ei ole enää pantakarhua. In Moilanen, A., Wilson, K.A., Puruvesi, (ed. Heimo Paakkinen), 29.9. & Possingham, H.P. Sundell, J. Interview ofn brown bear research. In (in press, scheduled Viiden kunnan sanomat, (ed. R. Vesterinen). for 5/2009) Spatial Sundell, J. Interviews on brown bear research. In. Conservation Prioritization: YLE Etelä-Savon radio, March 19th and April 9th. Quantitative methods Sundell, J. Karhun ohi tietämättä. In Koti-Kajaani, (ed. and computational tools. Mari Tikkunen), 6.9. Oxford University Press, Oxford. Sundell, J. Nähtäisiinpä karhu. In Metsälehti Makasiini, (ed. Jere Malinen), 3/2008. Sundell, J. Otso on epätoivottu vastaantulija. In Articles, interviews and programs in Länsi-Savo, (ed. Jarno Liski), 30.6. public media Sundell, J. Päin karhua! In Metsästys ja kalastus, (ed. Jere Malinen), “Luonnossa” suppl. 6/2008. Sundell, J. Voles! Help us! (in Finnish: Myyriä! Tulkaa Hanski, I. Butterfly effects. In HUB - Helsinki apuun!). In Suomen Luonto, 7/2008. University Bulletin, (ed. Päivi Lehtinen), 1/08, pp. van Nouhuys, S. Insect Behaviour: Learning for the 10-13. Future wasp, which oviposits in butterfly eggs. In Hanski, I. was interviewed on several newspapers Current Biology, (ed. TS Collet), 18, pp. R131-R134. and magazines about his work in the field van Nouhuys, S. A story about a wasp, a caterpillar of ecology. e.g. Helsingin Sanomat monthly and a fungus ends with more dominating female magazine. wasps. In Cornell Chronicle, (ed. K. Ramanujan), Kaartinen, R. Äkämäpistiäisiä etsimässä. In 23 Jan. Luontoretki. YLE Radio Suomi, 21.9. Wheat, C.W. & Vera, J.C. Sequencing goes 454 Ovaskainen, O. How to draw the boundaries of a and takes large-scale genomics into the wild. habitat. In HUB - Helsinki University Bulletin, (ed. In Molecular Ecology News and Views, (ed. H. Juha Merimaa), 1/08, pp. 44-45. Ellegren), 17, pp. 1629-1635. Ovaskainen, O. Starttirahaa tutkijoille. In Yliopisto, (ed. Tiedeuutinen), 2/2008. Roslin, T. Interview on dung beetle ecology. In Kvanthopp (ed. M. Rosenlund). YLE Radio Vega, Honours, awards and memberships October 22. Roslin, T. Interview on dung beetle ecology. In Mikko Frilander worked as an external expert in Ajantasa. YLE Radio Suomi, June 6. Estonian Research council. Roslin, T. Kiinnostavat lantaläjät. In Ympäristö, Ilkka Hanski was awarded Lauri Jäntti Foundation’s 4/2008. nonfiction literature prize for the book Viestejä Roslin, T. Kittilän lantakuoriaisia lähetetään etelään. Saarilta. In Kittilälehti, 25/2008. Heini Kujala was a member of Societas Biologica Roslin, T. Kumman huvittava tiede. In Matkaan, Fennica Vanamo and Society for Conservation 8/2008. Biology. Roslin, T. Lantakuoriaisia etsimässä. In Tiede, 7/2008. Tomas Roslin has been a member of the board of Roslin, T. Lehmän lantakasa tarjoaa elämän Societas Entomologica Helsingforsiensis. kymmenille lantakuoriaislajeille. In Maaseudun Janne Sundell was a member of editorial boards of Tulevaisuus, (ed. Markus Mykkänen), 18.6. Oecologia and Ecological Research. Roslin, T. Lehmänläjä kuhisee kuoriaiselämää. In Hämeen Sanomat, (ed. Pirkko-Liisa Kastari), 16.6. Roslin, T. Lehmänläjät tutkitaan tänä kesänä Turusta Inariin. In Kaleva, (ed. Pekka Rahko), 8.6. Roslin, T. Miksi kannattaa elää kengurun perskarvoissa. In Alasatakunta, (ed. Tuomo

46 Lehtomäki, J. “Conservation prioritization: Finnish Conferences and seminars boreal forests as an example”. Biodiversity in For- est Ecosystems and Landscapes (IUFRO interna- Arponen, A. “Yhteisömallinnuksen menetelmiä tional conference), August 5-8th, Poster, Kamloops, suojelualuesuunnittelussa (Community modelling Canada. methods in conservation planning)”. Finnish Envi- Lehtomäki, J. ”Paikkatietoaineistojen hyödyntäminen ronment Institute: Luonnonsuojelubiologian päivä, suojelualueverkon kokonaisvaltaisessa suunnitel- January 22th, Talk, Helsinki. lussa – in Finnish”. Ympäristöhallinnon metsäpäivät, Frilander, M.J. “A walk on the wild side: Molecular November 3-4th, Talk, Lammi. biology meets ecology”. Biocentrum Helsinki Lehtomäki, J. “Suojelualueiden ekologinen arvot- meeting, Jan 17th, Talk, Helsinki. taminen: Zonation – in Finnish”. Metsähallituksen Frilander, M.J. “Ecological genomics of Melitae cinxia luonnonsuojelupäivät, May 14th, Talk, Kuusamo. butterfly”. High Throughput Biology, June 12-13th, Lehtomäki, J. ”Uusi lähestymistapa metsiensuojeluun Talk, Helsinki. ja metsien hyötykäytön suunnitteluun - in Finnish”. Frilander, M.J. RNA in Biology, Biotechnology, and Luonnonsuojelubiologian päivä (SYKE), January Medicine, October 24-26th, Meeting organizer, 22th, Talk, Helsinki. Helsinki. Mitikka, V. ”Regional patterns in the expansion of the Hanski, I. Workshops on coupling between ecological map butterfly (Araschnia levana).” Future of butter- and evolutionary dynamics, January and Septem- flies in Europe, April 17-19th. Poster, Wageningen, ber, Workshop, Silwood Park, UK. The Netherlands. Hanski, I. Annual meeting of the Royal Entomological Niitepõld, K. “Melitaea cinxia: shaken not stirred. Society, September, Plenary talk, Plymouth, UK. Metabolic rate predicts mobility”, May, Talk, Catholic Hanski, I. University of Århus, September, Invited University of Louvain, Belgium. seminar, Århus, Denmark. Niitepõld, K. “Melitaea cinxia: shaken not stirred. Hanski, I. Conference on the conservation of boreal Studying metabolic rate and correlated traits”, May, forests, October, Invited speaker, Pedrozavodsk, Talk, University of Leiden, The Netherlands. Russia. Niitepõld, K. “Tracking butterfly flight with harmonic Hanski, I. Southwood Lecture in the University of radar: Linking genotype, phenotype and perfor- Oxford, October, Lecture, Oxford, UK. mance”. International Congress of Entomology, July, Hanski, I. Lecture in the University of Antananarivo, Talk, Durban, South Africa. November, Lecture, Antananarivo, Madagascar. Norros, V. “Measuring and modelling dispersal in Hanski, I. Centenary symposium on “Finland’s chang- wood-decaying fungi”. Effects of forestry on sap- ing nature”. Finnish Academy of Science and Let- roxylic insects and fungi, Organized by Norwegian ters, April 16th, Organizer, Helsinki. Forestry Association (NORSKOG) and the Norwe- Harrison, P.J. “Spatial structure and population dy- gian Institute for Nature Research (NINA). Work- namics in fragmented landscapes”. International shop, Vindfjelltunet Guesthouse, Norway. Statistical Ecology Conference, July 9-11th, Talk, St Ovaskainen, O. “Forward and inverse approaches in Andrews, Scotland. mathematical biology” and “The application process Hottola, J. “Talousmetsien luonnonhoidon ja va- for the ERC-starting grant”. University of Tarto, April paaehtoisten suojelukeinojen merkitys kääpien 15th, Talks, Tarto, Estonia. monimuotoisuudelle”. Societas pro Fauna et Flora Ovaskainen, O. “High throughput sequencing of Fennican symposium Näkyykö metsä puilta? - Ser environmental samples - what is the probability that man skogen för bara träd?, November 21st, Invited your BLAST result is the correct species?” Nordforsk talk, Helsinki. network meeting, September 28-29th, Talk, Tallinn, Kaartinen, R. Quantitative food webs in space: Estonia. herbivore-parasitoid communities on pedunculate Ovaskainen, O. “Kuka maksaa sukupuuttovelan oak Quercus robur. International Conference in En- Suomen luonnossa”. Muuttuva Suomen luonto tomology, July 6-12th, Talk, Durban, South Africa. -symposium, April 16th. Talk, Helsinki. Kujala, H. 22nd Annual meeting of the Society of Ovaskainen, O. “Living – or dying? – in fragmented Conservation Biology, July 13-17th, Talk, Chatta- environments”. Seminar on Current Environmental nooga, Tennessee, USA. Issues. Talk, University of Joensuu, Joensuu. Kujala, H. 2nd meeting of the NCEAS Connectivity Ovaskainen, O. “Modeling movement data with Working Group, July 17-19th, Talk, Chattanooga, Bayesian state-space approaches” The International Tennessee, USA. Statistical Ecology Conference, July 9-11th, Talk, St. Kujala, H. Science meeting at the Finnish Forest and Andrews, UK. Park Services (Metsähallitus), March 10th, Talk, Ovaskainen, O. Participated a UK Popnet meeting Vantaa. Bayesian distribution models: dynamics, processes and projections, September 7-11th, York, UK.

47 Roslin, T. SEBDEM symposium, April 1-2nd. External It was a collaboration with the International evaluator and plenary speaker, University of Turku. Mammoth Committee and the “Institute for Sundell, J. “Bears and humans: in Finnish”. Meeting Biological Problems of Cryolithozone” in Yakutsk. of Game Management District of Central-Finland. Heini Kujala visited Biodiversity and Global Talk, Keuruu, Finland. Change Lab in the Museum of Natural Sciences Sundell, J. “Behaviour of the brown bear in con- SCIC,Madrid, Spain. 1.–29.4. tacts with humans: in Finnish”. Public meeting of Riikka Kaartinen did a collaborative work with Southern Savo Game Management District. Talk, Graham Stone in the University of Edinburgh, Savonlinna. Edinburgh, Scotland. 26.10.–20.12. Sundell, J. “Behaviour of the brown bear in contacts with humans: in Finnish”. Conservation Biology Field work for dung beetle project in Madagascar: Day. Talk, Finnish Environment Institute, Helsinki. Mirja Miinala 15.11.–13.12. Tack, A. “The role of competition in structuring an Ilkka Hanski 15.11.– 3.12. insect metacommunity on oak (Quercus robur)”. Heidi Viljanen 7.1.–21.2. ICE conference 2008, July 6-12th. Talk, Durban, South Africa. van Nouhuys, S. Symposium on- Genetic and Ge- Visitors nomic Approaches for Parasitoid Behavioural Ecol- ogy, June 8th. Invited speaker, Scotland UK. Prof. Tom Collett and Andy Philippides, University van Nouhuys, S. & Kraft., T. “Complex pattern of indi- of Sussex, UK, collaboration with Saskya van rect interaction between butterflies due to shared Nouhuys, 1-11.07. pupal parasitoids”. Annual meeting of the Entomo- Dr. Stephen Cornell, Leeds University, UK, logical Society of America, November 8th. Talk. collaboration with Otso Ovaskainen, several visits. Wheat, C.W. “Expression phenotypes are sorted by Prof. Elizabeth Crone, The University of Montana, ecological dynamics”. Society for Molecular Biol- USA, collaborative work with Otso Ovaskainen, ogy and Evolution meeting, June 5-8th. Poster, 15.8.07–31.7.08. Barcelona, Spain. Prof. Charles Godfray, University of Oxford, UK, MRG Wheat, C.W. “Functional genomics of dispersal in the Center of Excellence Scientific Advisory Board wild”. Gordon Conference on Molecular Evolution, meeting, 26.–29.3. February 3-8th. Poster, Ventura, California, USA. Dr. Jenny Hodgson, Leeds University, UK, Wheat, C.W. “Functional Genomics of Dispersal: collaboration with Otso Ovaskainen, several visits. developing and using molecular tools in an eco- Dr. Laura Jones, Cornell University, collaborative logical model system”. Department of Biology, Uni- work with Saskya van Nouhuys, 31.7.–11.8.. versity of California, February 8th. Talk, Riverside, Tom Kraft, Cornell University, work with Saskya van Ventura, California, USA. Nouhuys, 17.5.–15.7. Wheat, C.W. “Metapopulation processes sort expres- Prof. Jim Marden and MsC Cristobal Vera, Penn State sion phenotypes”. Facultat de Veterinaria, Uni- University, PA, USA, collaborative work with Ilkka versitat Autonoma de Barcelona, June 10th. Talk, Hanski and Mikko Frilander, 16.–23.10. Barcelona, Spain. Prof. Michael Nachman, Universiyt of Arizona, USA, Viljanen, H. “Native and introduced dung beetles MRG Center of Excellence Scientific Advisory (Scarabaeidae: Aphodidae) using cattle dung in Board meeting, 26.–29.3. Madagascar”. XXIII International Congress of Ento- MSc Sandra Nogue, Universitat Autònoma de mology, July 6-12th. Talk, Durban, South Africa. Barcelona, Spain, 10.9.–21.12. Zheng, C. “Hierarchical metapopulation dynam- Prof. Reed F. Noss, University of Central Florida, ics of two aphid species on a shared host plant”. USA, MRG Center of Excellence Scientific Advisory International Statistical Ecology Conference, July Board meeting, 26.–29.3. 9–11th, Talk, University of St Andrews, St Andrews, MSc Carolina Reigada, Universidade Estadual UK. Paulista, São Paulo, Brazil, work with Saskya van Nouhuys and Ilkka Hanski, 7.1.–11.9. Prof. Derek Roff, Riverside, CA, USA, visiting the Work abroad group, 16.4. MSc Esther Sebastian Gonzales, Universidad Miguel Hernández, Alicante, Spain, 2.9.–3.11. Olivier Gilg did field work in Central Siberia, July– Prof. John N. Thompson, University of California at August. It concentrated mainly to inventor birds, Santa Cruz, US, visited Anna-Liisa Laine, 1.–10.8. mammals and vascular plants along the Anabar lower river (from the tree line to the Arctic Ocean).

48 Teaching

Spatial Population Ecology have had for the last two years, we supplement traditional lectures with a series of discussion groups leading up to an informal seminar. These discussion Uppsala University, Sweden, 1 week course in groups offer student the challenge of applying May the information they have learnt to real world conservation issues and of evaluating the relevance and scope of existing scientific information when it Post graduate workshop: Metacommunity Ecology comes to applied management issues – including Instructor: Saskya van Nouhuys the extent to which scientific information is actually used when real decisions are made.

Cornell University, USA, 9.-12.8. The students presented the results of these working Undergraduate course: Insect Biology groups during a two day seminar in November. To Instructor: Saskya van Nouhuys bring in the viewpoint of those working on applied conservation issues, we invited two external guests to the seminar: Sini Harkki (the coordinator of University of Helsinki, 6.-10.10. forest issues for Greenpeace); and Lauri Saaristo (a conservation advisor for TAPIO). During the seminar Undergraduate course: Ecology of small mammals the students gave two presentations per group Responsible teacher: Janne Sundell (a detailed talk aimed at a scientific audience and a shorter one aimed at a more general audience akin to an Environmental Impacts Assessment Conservation Biology in Fragmented meeting). The seminar was a great success, with all participants engaging in vigorous discussion and Landscapes participating in purpose invented games and role playing. University of Helsinki 29.9. - 10.12. Overall, we find it an exciting challenge to use our own research results to illustrate fundamental Coordinators: Phil Harrison and Tomas Roslin ecological concepts to the students. Our teaching unit also functions as a uniting factor, forcing us Since the year 2000, the Metapopulation Research to relate all the work conducted in our group to a Group has annually taught a course on Conservation uniform framework. Biology in Fragmented Landscapes (7 ECTS compulsory + 2 optional). The course is part of the international Boreal Biota and Ecology program, but Finnish students are also encouraged to participate. In 2007, we had 11 Finnish students and 15 Irish students. Our objective is to unite all MRG members in a joint venture, to compile a coherent teaching unit that takes advantage of our own research, and to provide group members with essential teaching experience. As in all our work, we blend theory and modelling with empirical case studies, and try to offer the students our particular view on how habitat fragmentation affects key aspects of population biology. Students are also given the choice of earning two extra ECTS by reading books produced in the group - this year The Shrinking World by Ilkka Hanski. © Elmira Zaynagutdinova

In 2008, the course was coordinated by Phil Harrison & Tomas Roslin. Following on from an initiative we

49 Budget

Funding source EUR

Academy of Finland Centre of Excellence funding Metapopulation Research Group 413 150 Academy professor's research costs Ilkka Hanski: Research on interaction of spatial and evolutionary dynamics 177 150 Academy research fellow's research costs Atte Moilanen: Reserve network design: ecological and economical factors 6 860 Otso Ovaskainen: Mathematical theory for population dynamics in fragmented 118 210 landscapes Tomas Roslin: Metacommunity dynamics of oak-associated organisms 126 030 Postdoctoral projects Phil Harrison: Spatial structure and population dynamics in fragmented landscapes: 60 000 combining diverse data sets for improved inference + 1 Academy professor post and 3 Academy research fellow posts

University of Helsinki Centre of Excellence funding 295 500 Ilkka Hanski & Mikko Frilander: Biocentrum Helsinki 45 000 + 1 PhD student post and 1 Laboratory technician post

European Union Mar Cabeza: Minimization of and adaptation to climate change: Impacts on biodiver- 28 270 sity Otso Ovaskainen: Spatial Ecology: Bringing mathematical theory and data together 125 120

Foundations Ella and Georg Ehrnrooth Foundation + 1 personal grant for a post doctoral researcher Kone foundation Tomas Roslin: Community structure and function in fragmented landscapes 6 810 + 1 personal grant for a post doctoral researcher Maj and Tor Nessling Foundation Mar Cabeza: Conservation area planning and climate change 20 225 Oskar Öflunds Stifelse Janne Sundell: Antipredatory behaviour of the brown bear 2 500

50 National Science Foundation (USA) Jim Marden & Ilkka Hanski: Functional Genomics of a Metapopulation: An Individual 27 500 Based Understanding of Population Dynamics, Life History Traits and Geneticecha- nisms in Time and Space + personal grant for a postdoctoral researcher Svenska Kulturfonden Tomas Roslin: Community structure and function 2 000

LUOVA graduate school 3 PhD student posts

TOTAL 1 455 325 + research posts/personal grants

All over 10 000 eur grants from Academy of Finland, foundations and EU include 15-20% overhead The given sums for EU and Foundations funding are estimates for year 2008 if the funding is appropriated for several years.

Sources of Metapopulation Research Group funding in 2008.

51 Prospects for the year 2009

e will start, in the beginning of 2009, docs, and there will be students and post docs with the second 3-yr period of our current expertise that we are currently lacking. There will be WCentre-of-Excellence funding cycle. The a big increase in molecular and genomics work and CoE program has been critically important for us. extensions of our current strength in mathematical There is no doubt that without the CoE core funding modelling and application of various numerical the breadth and the depth of the research in MRG methods. would not be what it is today. Though we will never know for sure, the CoE funding has helped us to One could even question whether we are moving secure other sources of funding, including the too fast to areas that are new to us. Is there a current membership in Biocentrum Helsinki, a big danger that we fail to capitalize on our past and Biocomplexity grant from US NSF together with Jim current strengths and attempt to initiate research Marden (2005-08), and two recent big grants from for which we do not have sufficient knowledge the European Research Council. and resources? There may be such a danger, but the answer is not to stick to what we have done Every year brings something new, but 2009 will in the past but to be aware of the challenges in be exceptional, perhaps the year of the biggest entering into new type of research. And we have changes so far in the entire history of our research to constantly keep in mind our core mission: to group. First, perhaps because of the funding cycle develop better understanding of population, and the start of the current CoE funding 3 years evolutionary, and conservation biology in the ago, no PhD student finished in our group in 2008. spatial context. The use of new genomics tools is This will be more than compensated for by the potentially a very powerful approach, and we are in exceptionally large number of students finishing in a good position to make important contributions 2009. To compensate for the losses, we will have an to integrated biology, bringing together knowledge extra large number of new students and post docs from molecules to landscapes. joining us in 2009. Metapopulation Research Group has been an Not only is the number of new people going to be exciting place to do research in the past. Its going to exceptionally large, their composition will also be be even more exciting in 2009! exceptional. There will be several new senior post Ilkka Hanski