Annual Report 2003 Annual Report 2003
Metapopulation Research Group
Department of Ecology and Systematics
University of Helsinki
Edited by Tapio Gustafsson Contact information
Address: Metapopulation Research Group Department of Biological and Environmental Sciences PO Box 65 (Viikinkaari 1) 00014 University of Helsinki FINLAND
Phone: +358 9 1911 (Exchange) Fax: +358 9 191 57694
E-mail: [email protected]
Web site
Metapopulation Research Group (UH), http://www.helsinki.fi/science/metapop/
MRG-logo designed by Gergely Várkonyi
Cover photo: The Namorona River runs throuh the rainforest of the Ranomafana National Park in Madagascar. Photo by Tapio Gustafsson Contents
Preface...... 6
Brief history and overview of the MRG...... 7
Projects in the MRG ...... 10 Ecology, genetics, and evolution of metapopulations ...... 10 On the wings of checkerspots: A model system for population biology ...... 11 Metapopulation theory ...... 12 Population genetics theory ...... 13 Metapopulation dynamics and reserve network design ...... 14 Spatial ecology of arboreal insects ...... 15 Metapopulation biology of the Glanville fritillary butterfly ...... 16 Biology of the parasitoids of the Glanville fritillary...... 17 Demography and genetic structure of shrew populations on small islands...... 18 Ecology of small rodents and their predators ...... 19 Spatial ecology and human interactions of the brown bear ...... 20 Population biology of trembling aspen ...... 21 Gene flow in aspen ...... 22 Spatial ecology of plant-pathogen interactions ...... 23 Biology of an old adaptive radiation: Evolutionary community ecology of the beetle tribes Canthonini and Helictopleurini in Madagascar...... 24 Influence of climate change on the distribution and abundance of species ...... 25 Biodiversity in boreal forests ...... 26
MRG personnel ...... 27
Laboratory facilities ...... 61 Field sites ...... 63 Åland Islands...... 63 Kuhmo ...... 63 Greenland ...... 65 Wattkast ...... 66 Madagascar: Ranomafana National Park ...... 67
Synopsis of the year 2003 ...... 69 Budget ...... 69 Publications ...... 70 Theses ...... 75 External visits ...... 80 Seminars, lectures, and talks ...... 81 TV, radio, and newspapers ...... 85 Teaching and courses ...... 87 Honors and awards ...... 89 Council memberships ...... 89 Meetings organized by the MRG ...... 90 Visitors to the MRG ...... 90
Prospects for the year 2004 ...... 92 5
Occupied (filled) and empty (open) habitat patches suitable for the Glanville fritillary in the Åland Islands in the autumn 2003.
MRG students and researchers searhing for Melitaea cinxia larvae during our Annual Meeting in the Åland Islands in May. Photo by Evgeniy Meyke. 6 - Preface Preface
The past year was our first full year in the new campus. It must be a sign of things going pretty well here in Viikki that one does not hear any more comments about what went before in the old building in downtown. Office space continues to be a problem, but this is what we expected - and I am glad to report that everybody did have a desk in 2003. New exciting projects were started in 2003. My own memories from June relate to the hugely successful experiments with the Glanville fritillary that were accomplished in the large cage that was erected in Åland in the spring. I would be very surprised if this experimental set-up would not turn out to be another hall mark of the long-term research in Åland; the opportunities for truly novel research are definitely there. The new project on tropical forest dung beetles in Madagascar was officially launched with a MRG expedition landing in Antananarivo in November. This was the real highlight of the year for many of us. I was busy until the very end of the year with the two edited volumes that are now, at last, in press and will appear in early 2004, the Ehrlich & Hanski volume on the biology of checkerspots and the new Hanski & Gaggiotti metapopulation volume. There were disappointments in 2003, too. The checkerspot project in China had to be cancelled at a short notice because of the SARS epidemics. The 6th Framework Programme of the EU has not much to offer to European population biologists and, sadly, it appears, population biologists have not much to offer to the EU. The structure of our research group continued to change in 2003. I suppose that the temporal pattern in the relative numbers of graduate students, post docs, and senior researchers that is evident in the graph on p. 7 is rather typical for the first 10 years of most research groups that were started with just one researcher. First a rapid increase in the numbers of graduate students; then a period of disproportionate numbers of post docs; followed by an increase in senior researchers, reduction in post docs, and stabilization of the numbers of graduate students. The MRG has now experienced all these stages, as well as the departure of the first senior researchers. Oscar Gaggiotti moved to a professorship in Grenoble and Marko Nieminen to a full-time position in his consulting company. Thanks to Oscar and Marko for all your contributions to the MRG. Two MRG researchers have progressed to the level of senior researcher, Tomas Roslin and Saskya van Nouhuys. Three students completed their PhD in 2003, Mar Cabeza-Jaimejuan, Gergely Várkonyi, and Sari Haikola, while Johanna Gripenberg, Mervi Koskela, Varpu Mitikka, and Astrid van Teeffelen started. Finally, the ranks of our junior members increased less than in the past years, but one event was nonetheless recorded also in this front. Our warmest congratulations to Tapio and Riitta (girl).
Ilkka Hanski History and overview - 7 Brief history and overview of the MRG
Professor Ilkka Hanski has worked on spatially structured populations since the late 1970’s. The early work dealt primarily with small-scale spatial population structure, but since the early 1980’s the focus shifted to larger spatial scales and to metapopulation dynamics in the sense of assemblages of discrete local populations connected by migration. In 1989, Hanski organized the first international meeting on metapopulation dynamics together with Professor Michael Gilpin (San Diego, UC), which resulted in the first edited volume on the subject (Gilpin & Hanski, 1991, Metapopulation Dynamics: Empirical and Theoretical Investigations, Academic Press, London). This meeting led to the conception of the MRG. Ongoing collaboration with Professor Mats Gyllenberg (Department of Mathematics, Turku, Finland) started in 1990, the long-term field project on the Glanville fritillary butterfly in the Åland Islands was started in 1991, and the first post graduate students and post docs were accepted in the MRG in 1992 and 1993, respectively. The national research council (the Academy of Finland) nominated the MRG as one of the Centres of Excellence in Research in 2000-2005 (www.aka.fi).
The figure below illustrates the growth of the MRG since 1992. 8 - History and overview Currently there are 15 post graduate students, 3 post docs and 6 senior researchers, and the technical staff includes 4 persons. The senior researchers have expertise from ecology (Ilkka Hanski, Tomas Roslin), butterfly biology (Marko Nieminen), and parasitoid biology (Saskya van Nouhuys) to theoretical population genetics (Oscar Gaggiotti) to computing science (Atte Moilanen) and mathematics (Otso Ovaskainen). Personnel in the MRG represents 11 different nationalities from 4 continents.
Home institution
The MRG is the largest research group in the Division of Population Biology (DPB), Department of Ecology and Systematics at the University of Helsinki.The permanent staff of the DPB consists of 3 professors and 9 other academic staff. Despite its small size the DPB with several active research groups and researchers has become an internationally recognized centre of population and evolutionary biology. There are currently 35 post graduate students and 28 post docs in the DPB, many of whom come from elsewhere in Europe and from further away. The MRG has had an influence on the general direction of research in the DPB, which is now to a large extent focused on spatial ecology and population biology. Since 1997, the DPB has run a joint research programme called Spatial Ecology (see www.helsinki.fi/ml/ekol/spatial_ecology.html).
Scientific standing
The MRG has established a solid reputation as a leading research group internationally in metapopulation biology. The two most visible achievements of the MRG are the development of effective modelling approaches to the dynamics of metapopulations living in highly fragmented landscapes and a large-scale empirical research project on the Glanville fritillary butterfly (Melitaea cinxia). The latter started as an ecology project but has by now expanded to cover metapopulation genetics and evolutionary biology as well. The Glanville fritillary in the Åland Islands has effectively become a model system for metapopulation biology. It has stimulated new theory and it has allowed us to test many theoretical predictions; the Glanville fritillary system functions as an important interface between theory and empirical research in metapopulation biology. History and overview - 9
Strategic goals of the Metapopulation Research Group
* To strengthen our position as the leading research group in metapopulation biology worldwide
* To build up on our current strength in combining modelling with empirical studies
* To integrate genetic and evolutionary studies into the existing strong ecological framework and thereby to promote a comprehensive approach to metapopulation biology
* To facilitate the application of metapopulation biology in landscape-level environmental planning and management and conservation of populations.
The brown bear (Ursus arctos) is one of the new study species in the MRG. Photo by Jonna Katajisto. 10 - Projects Projects
Ecology, genetics, and evolution of metapopulations
Researchers: Ilkka Hanski and Oscar Gaggiotti
This volume edited by Ilkka Hanski and Oscar Gaggiotti presents a current overview of metapopulation biology, integrating over ecological, genetical and evolutionary consequences of habitat fragmentation. The volume consists of 23 chapters written by leading authors in the field, covering both recent theoretical advances and some of the best empirical studies applying the metapopulation approach. The MRG authors include Ilkka Hanski, Oscar Gaggiotti, Otso Ovaskainen, Atte Moilanen, and Mar Cabeza. The manuscript was completed during the year 2003, and the book will be published by Academic Press in February 2004.
Key publication:
Hanski, I. and Gaggiotti, O. (eds.). 2004. Ecology, genetics, and evolution of metapopulations. Academic Press, February 2004.
Collaborators:
32 researchers working in different fields of metapopulation biology. Projects - 11 On the wings of checkerspots: A model system for population biology
Researchers: Ilkka Hanski, Marko Nieminen, and Saskya van Nouhuys Technician: Mika Siljander
This is a volume on the population biology of checkerspot butterflies edited by Paul R. Ehrlich and Ilkka Hanski. We have aimed at producing a synthesis of the long- term studies by Ehrlich (Stanford) and his group on Euphydryas editha and other North American species, and of our studies on Melitaea cinxia and other European species. Chapters cover topics such as taxonomic and ecological background, population structure and dynamics of E. editha and M. cinxia, reproductive biology, biology of oviposition preference, larval biology, natural enemies, dispersal behaviour, population genetics, and conservation implications of our studies. The current and ex-MRG authors include Ilkka Hanski, Niklas Wahlberg, Marko Nieminen, Mika Siljander, Mikko Kuussaari, Saskya van Nouhuys, and Ilik Saccheri. The manuscript was completed in the spring 2003. The book will be published by Oxford University Press in February 2004.
Key publication:
Ehrlich, P.R. and Hanski, I. (eds.). 2004. On the wings of checkerspots: A model system for population biology. Oxford University Press, February 2004.
Collaborators:
Prof. Paul R. Ehrlich and his research group, Stanford, US. 12 - Projects Metapopulation theory
Researchers: Otso Ovaskainen and Ilkka Hanski
Our leading theme in metapopulation theory has been in assessing the conditions under which a given species is or is not expected to persist in a given fragmented landscape. Much of our work has been based on stochastic patch occupancy models (SPOMs), which characterize a landscape as a network of habitat patches, each patch being classified simply as occupied or empty. We have combined SPOMs with assumptions about how the landscape structure will influence population extinctions and recolonizations, which leads to what we have dubbed spatially realistic metapopulation theory (SMT; Hanski & Ovaskainen 2003). To better understand the dispersal phase in metapopulation dynamics, we supplemented the diffusion approximations of correlated random walks with boundary conditions corresponding to edge-mediated behaviour (Ovaskainen & Cornell 2003). We utilized the theory of stochastic processes (e.g. first-passage processes) and the theory of partial differential equations (e.g. adjoint operators and the finite-element method) to analyze and parameterize models that quantify movement behaviour in real heterogeneous landscapes (Ovaskainen 2004). We developed individual-based modelling approaches to address evolutionary questions such as the evolution of host plant oviposition preference and evolution of migration rate. These studies combine theory and empirical work in the context of the Glanville fritillary project (see page 16).
Key publications:
Hanski, I. and Ovaskainen, O. 2003. Metapopulation theory for fragmented landscapes. Theoretical Population Biology, 64, 119-127.
Ovaskainen, O. and Cornell, S. 2003. Biased movement at a boundary and conditional occupancy times for diffusion processes. Journal of Applied Probability 40, 557-580.
Ovaskainen, O. 2004. Habitat-specific movement parameters estimated using mark– recapture data and a diffusion model. Ecology 85, 242-257.
Collaborators:
Dr. Stephen Cornell, Cambridge University, UK Prof. Kevin Higgins, South Carolina, US Dr. Mikko Heino, Trömsö, Norway Projects - 13 Population genetics theory
Researcher: Oscar Gaggiotti
The objective is to develop new population genetic theory and statistical genetic methods aimed at incorporating the ecological complexities of natural populations. This line of research will lead to a more thorough integration of the fields of demography, population ecology and population genetics. Statistical modelling: Recently developed computer intensive methods open the field to the development of statistical genetic methods that are more appropriate for the study of metapopulations. At the moment there are two lines of research being carried out in this area: (1) Statistical genetic methods for the study of metapopulation processes: I have recently completed the development of a method to study colonisation processes. The results will be published in Molecular Ecology at the end of this year. This line of research will be continued with the development of statistical genetic methods to study migration in metapopulations and a method for the simultaneous estimation of Wright’s inbreeding coefficients. (2) Theoretical modelling: Classical population genetics theory has largely ignored the effect of age- and stage-structure on the genetics of fragmented populations. The lines of research in this area include the effect of age-and stage-structure: (a) on effective population size, and (b) on spatial genetic structure. At the moment I’m applying these ideas to populations of the macroalgae Gracilaria gracilis. In the near future I will developed similar models for metapopulations of the grey seal and a subterranean rodent (Ctenomys australis).
Key publications:
Gaggiotti, O.E., Jones, F., Lee, W.M., Amos, W., Harwood, J. and Nichols, R.A. 2002. Patterns of colonization in a grey seal metapopulation. Nature 416, 424-427.
Gaggiotti O.E. 2004. Multilocus Genotype Methods for the study of Metapopulation Processes, in Metapopulation Ecology, Genetics and Evolution, Hanski, I. and Gaggiotti, O.E. (eds.), Academic Press.
Gaggiotti, O.E., Brooks, S.J., Amos, W. and Harwood, J. 2004. Combining demographic, environmental and genetic data to test hypothesis about colonization events in metapopulations. Molecular Ecology (in press). 14 - Projects Metapopulation dynamics and reserve network design
Researcher: Atte Moilanen Post doc: Mar Cabeza Post graduate student: Astrid van Teeffelen Undergraduate students: Anni Arponen and Joona Lehtomäki
Reserve selection (RS) concerns the planning of reserve networks that would represent well the full spectrum of biodiversity in the region. Computational RS methods aim at finding cost-effective and optimal solutions to RS problems. Until a few years ago, RS methods didn´t include any notion of spatial (meta)population dynamics and consequent inevitable changes in species distributions (see Cabeza and Moilanen 2001 for review). The methods that are being developed in the RS project aim at the inclusion of spatial population dynamics into computational RS methods so that long- term persistence of biodiversity is maximized. In this project we have so far developed RS methods that consider connectivity, spatial dynamics and long-term persistence in RS (Moilanen and Cabeza 2002, Cabeza 2003, Cabeza et al. 2004). Currently we are extending these methods methods by (i) a stronger component of habitat modeling (AvT, AM), (ii) application to changing environments, e.g. in the context of climate change (MC, AM), (iii) improved optimization methods (AM), and (iv) by inclusion of uncertainty analysis (AM) and concepts such as threats and vulnerability (MC). Other related lines of research include application of the maximum coverage method (AA, AM) and reserve planning for powerline clearings (JL, AM).
Key publications:
Cabeza, M. and Moilanen, A. 2001. Design of reserve networks and the persistence of biodiversity. TREE 16:242-247.
Moilanen, A. and Cabeza, M. 2002. Single-species dynamic site selection. Ecological Applications 12:913-926.
Cabeza, M. 2003. Habitat loss and connectivity of reserve networks in probability approaches to reserve design. Ecology Letters 6: 665-672.
Collaborators:
Prof. Chris Thomas´ group, University of Leeds Prof. Mark Burgman´s group, University of Melbourne. Projects - 15 Spatial ecology of arboreal insects
Researcher: Tomas Roslin Post graduate student: Sofia Gripenberg
From the perspective of a host-specific herbivore, individual crowns of the host tree species represent islands of suitable habitat in a sea of unsuitable habitat. This project examines how the spatial distribution of such islands affects the interplay between the herbivore and its host. We strive to identify the role of spatial location in determining patterns of community composition, population dynamics and microevolution among monophagous insects on the English oak (Quercus robur). When modelling trees as islands, a crucial question emerges: how different are the islands from an insect’s perspective? The main achievement of this year was the completion of a chemoecological study initiated in 2001. Focusing on hydrolysable tannins and flavonoid glycosides, we quantified seasonal, inter- and intraindividual variation in the chemical content of oak leaves. A tree particularly rich in a given phenolic compound in the spring was often comparatively poor in this compound later in the summer. Combined with large variation in phenolic contents within trees, and in the performance of the moth Tischeria ekebladella, our results depict oaks as highly heterogeneous resources in both time and space. From a spatial perspective, the main accomplishment of year 2003 was the GPS-based mapping of more than 1,600 oak trees within an island approximately 5 km2 in size. This information has now been turned into a GIS data base. In the next few years, it will allow us to ask a broad range of questions on the role of spatial location in the biology of oak-associated herbivores.
Key publications:
Roslin, T., Salminen, J.-P., Vienola, M., Sinkkonen, J., Pihlaja, K., and Pulkkinen, P. Seasonal variation in the content of hydrolysable tannins and flavonoid glycosides in oak leaves. Submitted to Journal of Chemical Ecology.
Roslin, T., Gripenberg, S., Salminen, J-P., Vienola, M., O’Hara, R. B., Pihlaja, K. & Pulkkinen, P. Seeing the trees for the leaves – oaks as mosaics for a host-specific moth. Submitted to Oikos.
Collaborators:
Prof. Pertti Pulkkinen, Finnish Forest Research Institute, Finland Dr Juha-Pekka Salminen, University of Turku, Finland 16 - Projects Metapopulation biology of the Glanville fritillary butterfly
Researchers: Ilkka Hanski, Oscar Gaggiotti, Marko Nieminen, and Saskya van Nouhuys. Post graduate students: Sari Haikola, Marjo Saastamoinen, and Alia Sarhan Research assistants: Evgeniy Meyke Undergraduate students: Claudia Erälahti, Hanna Paulomäki, and Kata-Riina Valosaari.
This project was started with life history and mark-release-recapture studies in 1991. Since 1993, we have surveyed the entire Åland Islands twice a year for the presence and sizes of local populations in the around 4,200 small habitat patches. The major achievements in 2003 include highly successful experiments in a large (30 by 30 m) outdoor cage in Åland, in which we studied two experimental populations of around 200 butterflies. Butterflies were sampled for DNA and marked. We observed directly the vast majority of matings and ovipositions and will resolve the rest with the genetic markers. These data allow e.g. comparisons of life-time reproductive success of males and females originating from different kinds of populations. We have completed a study involving the [ATP]/[ADP] ratio of the flight muscles of butterflies. The results indicate that migration propensity of females is correlated with the [ATP]/[ADP] ratio. We have initiated new research on the Glanville fritillary in the Xinjiang province in China in collaboration with Prof. Guangchun Lei and his group in Beijing.
Key publications:
Hanski, I. and Heino, M. 2003. Metapopulation-level adaptation in insect host plant preference and extiction-colonization dynamics in heterogeneous landscapes. Theoretical Population Biology 64, 281-290.
Nieminen, M., Suomi, J., van Nouhuys, S., Sauri, P. and Riekkola, M.-L. 2003. Effect of iridoid glycoside content on oviposition host plant choice and parasitism in a specialist herbivore. Journal of Chemical Ecology 29, 823-844.
Collaborators:
Prof. Marja-Liisa Riekkola, University of Helsinki, Finland Prof. Michael Singer, University of Texas, USA Dr. Constantí Stefanescu, Butterfly Monitoring Scheme, Spain Dr. Johanna Suomi, University of Helsinki, Finland Prof. Jim Marden, Penn State, USA Prof. Guangchun Lei, Beijing, China Projects - 17 Biology of the parasitoids of the Glanville fritillary
Researchers: Saskya van Nouhuys and Ilkka Hanski Post graduate student: Maaria Kankare Undergraduate students: Johanna Ehrnsten, Riikka Kaartinen, and Eeva Punju
We work on the spatial dynamics and genetic structure of parasitoid populations, and the multitrophic interactions among parasitoids, their butterfly hosts and their host food plants. Over the years we have learned quite a bit about the natural history and spatial population dynamics of the primary parasitoids and hyperparasitoids hosted by the Glanville fritillary in the Åland islands. These data illustrate well the contrasting and sometimes complicated roles that dispersal and habitat fragmentation play in different host-parasitoid interactions, and in the competitive interactions among parasitoids. Our interests have expanded to parasitoid assemblages associated with Melitaeine butterflies throughout Europe, where both Melitaeine species richness, and inhabited landscape structure, vary. Parasitoids in the genus Cotesia are main characters in all of these assemblages, though their prevalence differs among communities. Recently we completed a molecular phylogeny of the Cotesia associated with Melitaeine butterflies, which revealed host associated cryptic species groups. We use microsatellite markers as tools to learn more about gene flow among Cotesia populations, and within and among the cryptic species.
Key publications: van Nouhuys, S. and Lei, G-C. 2004. Parasitoid and host metapopulation dynamics: the influence of temperature mediated phenological asynchrony. Journal of Animal Ecology, in press.
Kankare, M. and Shaw, M.R. 2004. Molecular phylogeny of Cotesia (Hymenoptera: Braconidae: Microgastrinae) parasitoids associated with checkerspot butterflies. Molecular Phylogenetics and Evolution, in press.
Collaborators:
Dr. Arjen Biere, Netherlands Institute of Ecology, The Netherlands Dr. Juan Corley, National Scientific Research Council of Argentina, Argentina Dr. Jeff Harvey, Netherlands Institute of Ecology, The Netherlands Dr. Mark Shaw, National museum of Scotland, Scotland Dr. Constantí Stefanescu, Butterfly Monitoring Scheme, Spain 18 - Projects Demography and genetic structure of shrew populations on small islands
Researchers: Ilkka Hanski and Gavin Hinten Undergraduate students: Aleksi Päivärinta, Anna Tuhti and Kaisa Välimäki
The shrew project is focused on genetic metapopulation dynamics in an otherwise well-studied system, where the spatial structure and the ecological dynamics of the metapopulation are known. More specifically, our aim is to integrate patterns of genetic diversity, measured with microsatellite markers, with ecological information from island and mainland metapopulations of three species of shrew, Sorex araneus, S. caecutiens, and S. minutus. We have samples from three regions: lake Sysmä and lake Koitere in eastern Finland, where many small populations on islands have been sampled, and a series of larger but more isolated islands in southwestern Finland. We have conducted field experiments to assess the strength of inbreeding depression by introducing pregnant female common shrews (S. araneus) from small islands (thought to be inbred) and mainland (outbred) to previously empty small islands. We will follow the reproductive success of the introduced shrews with the help of microsatellite markers. We have examined the relationship between genetics and fitness in intraspecific competition for resources. This was done by conducting neutral arena tests of intraspecific competition between individuals originating from island and mainland populations. The relatedness of individuals in the study populations will be assessed with microsatellite markers. The major achievements of the project in the past year include an individual- based simulation model, which incorporates a realistic description of the movement behaviour and demography of shrews as well as the geometry of the islands in lake Sysmä. With this model, we will compare the predicted and observed patterns in the genetic structure of the island populations. We have presently more than 10 highly polymorphic microsatellite markers for the common shrew (S. araneus) and are in the process of testing them on the other species. Around 1,000 common shrews have been genotyped and the material is currently being analysed.
Key publication:
Landry P.-A. 2004. Mainland-island metapopulation genetics of the common shrew (Sorex araneus). Proceedings of the second international colloquium on the biology of the Soricidae, in press.
Collaborators:
Dr. Pierre-Alexandre Landry, Montreal, Canada Projects - 19 Ecology of small rodents and their predators
Researchers: Olivier Gilg, Ilkka Hanski, and Janne Sundell Post graduate student: Paavo Hellstedt
The aim of the project is to obtain a better understanding of the role of predation in small rodent population dynamics by using both theoretical and empirical approaches. Empirical studies include large-scale spatial dynamics of voles, space use and habitat selection of stoats, factors affecting the survival of least weasels, and the numerical and functional responses of the predators of voles and the collared lemming. Experiments have been conducted in the laboratory, outdoor enclosures, and in the field. Field experiments have tested the role of mustelid predators in driving the vole cycle and the indirect effects of the presence of the predators on the behaviour and demography of voles. The most recent achievements of the project include empirical study and modelling of the impact of four species of predators on the dynamics of the collared lemming in eastern Greenland. The study involved detailed investigation of the predators’ total responses, which data were used to parameterise a predator-prey model. The major result was that the regular 4-yr cycle of the collared lemming in Greenland can be explained by predation only, without any density-dependent mechanisms in the prey population. According to our results, the lemming cycle is driven by predation by the stoat, the only resident specialist predator in the area, while the increase of the lemming population is stopped by density-dependent predation by the three other predators (arctic fox, snowy owl, and long-tailed skua) in the summer.
Key publications:
Gilg, O, Hanski, I. and Sittler, B. 2003. Cyclic dynamics in a simple vertebrate predator-prey community. Science 302, 866-868.
Sundell, J. 2003. Population dynamics of microtine rodents: an experimental test of the predation hypothesis. Oikos 101, 416-427.
Collaborators:
Heikki Henttonen, Finnish Forest Research Institute, Finland Erkki Korpimäki, University of Turku, Finland Benoît Sittler, University of Freiburg, Germany Hannu Ylönen, University of Jyväskylä, Finland 20 - Projects Spatial ecology and human interactions of the brown bear
Researchers: Atte Moilanen and Janne Sundell Post graduate students: Jonna Katajisto
In Finland and Scandinavia brown bear populations are expanding. Consequently conflicts between men and bears are likely to increase. In contrast, other European populations are relatively small and threatened. Habitat fragmentation and increasing human access to bear habitat together with illegal hunting pose the largest threats to bear populations. Clearly both landscape and human influence play a role in brown bear conservation and management. We have two subprojects to clarify these factors. Both have raised public interests resulting in magazine and television interviews. To assess bear population viability and prevent conflicts with humans it is necessary to understand how the spatial arrangement of habitats and human settlements affects the spatial distribution of bears. In the spatial population dynamics project (JK, AM) we have studied the effects of landscape structure and human activity on bear movements and home range establishment. Results so far show that bear movements and home range establishment are more limited by human influence than by natural factors related to habitat quality. Our goal is to construct a spatially explicit population model (SEPM) for predicting the future distribution of bears under different management scenarios. This work is done in collaboration with the Scandinavian Brown Bear Research Project, which provides us radiolocation data collected since 1984 from nearly 300 bear individuals. The other subproject investigates detailed behaviour of bears in human vicinity by using a new method of GPS-GSM tracking (JS). This method allows real time observations on bear activity and movements. Bear behaviour can be studied by conducting experiments in which bears will be approached in different habitats and in different manners. Specific behavioural responses, such as escaping speed and direction, possible hiding or aggressive behaviour, and reaction distances of bears will be obtained. Also possible changes in behaviour near the human settlement can be observed.
Collaborators:
Jon Swenson, Agricultural University of Norway Sven Brunberg, Scandinavian Brown Bear Project, Sweden Ilpo Kojola, Finnish Game and Fisheries Research Institute, Finland Thorsten Wiegand, UFZ Centre for Environmental Research Leipzig, Germany Eloy Revilla, Estación Biológica de Doñana, Spain Projects - 21 Population biology of trembling aspen
Researcher: Leena Suvanto Post graduate student: Tarja Salmi
This project was based on the finding that much research was done on species dependent on aspen, but not much was known about the population biology of the tree itself. As a result, the aspen project started in the beginning of year 2000. The main objectives of this project are to study the clonal structure and reproduction of aspen and the structure of aspen stands. Throughout this project a special emphasis has been put on the comparison of natural (old growth) and managed forests. One of the most important aspects of the study of clonal structure of aspen was the development of a reliable method for clone identification. This study has also looked at the amount and size of clones in the forests. Furthermore, the ages of the trees were determined in order to get a picture of the age distribution within and between clones. This study has been done by Leena Suvanto. The study of the structure of aspen stands at the landscape level is based on an earlier compiled data set on ca 30 000 spatially referenced mature aspen trees. In this work the amount of dead and living aspen wood in old growth and managed forests was determined. The spatial aggregation of coarse woody debris as well as the age distribution and the history of stand development were also studied. Tarja Salmi is responsible for this project. Aspen is capable of reproducing both asexually and sexually. Tarja Salmi has been studying the vegetative reproduction and the work connected to sexual reproduction has been done by both TS and LS.
Key publication:
Latva-Karjanmaa, T., Suvanto, L., Leinonen, K. and Rita, H. 2003. Emergence and survival of Populus tremula seedlings under varying moisture conditions. Can. J. For. Res. 33(11), 2081-2088.
Collaborators:
Juha Siitonen, Finnish Forest Research Institute, Finland. Kari Leinonen, Plant Production Inspection Centre, Finland. Hannu Rita, University of Helsinki, Finland. 22 - Projects
Gene flow in aspen
Researcher: Leena Suvanto Technician: Toshka Nyman
This project started in the beginning of year 2003 as a co-operation project with Finnish Forest Research Institute. It is dealing with the gene flow both within aspen as well as between aspen and hybrid aspen (Populus tremula x P. tremuloides). The work done in 2003 was aimed to clarify whether hybrid aspen could be a threat (in the form of gene flow) to native aspen. Hybrid aspen has been planted to Finland already from 1950’s, but the potential ecological or genetic hazards of these plantations have not been studied. We have studied the amount and survival of hybrid aspen seedlings, the sprouting ability of different aspen and hybrid aspen clones, crosses between aspen and hybrid aspen, and the detection of the origin of juveniles. The studies have been carried out in Southern Finland and Estonia. The results show that hybrid aspen can produce viable seeds and seedlings, but the survival of the seedlings is poor. There are differences in the sprouting capacity of different clones, but not between the two species. Fathers had no effect on the amount and viability of the seeds, but mothers and father-mother combination affected significantly both traits. Hybrid aspen produced more and more viable seeds than aspen and the amount and viability of the seeds was bigger, when the mother and father were of different species. The origin of the seedlings could not quite be determined with microsatellites, which was mainly due to lost mother genotypes (e.g. logging).
Key publication:
Suvanto, L., Stenvall, N., Vares, A. and Pulkkinen, P. 2004. Alien genes in forestry: hybrid aspen as a model species for alien gene flow. Rit Mogilsar Rannsöknastöovar Skograektar, in press.
Collaborators:
Doc. Pertti Pulkkinen, Finnish Forest Research Institute, Finland MSc Niina Stenvall, Finnish Forest Research Institute, Finland MSc Aivo Vares, Estonian Agricultural University, Estonia PhD Kari Leinonen, Plant Production Inspection Centre, Finland PhD Ari Pappinen, University of Helsinki, Finland Projects - 23 Spatial ecology of plant-pathogen interactions
Researcher: Ilkka Hanski Post graduate student: Anna-Liisa Laine Research assistant: Saila Kuokkanen
The dynamics of natural populations of plant pathogens are influenced by complex interactions both with the host plant and the environment, and these interactions vary both temporally and spatially. A full understanding of the dynamics of the pathogen requires studies at the metapopulation level where the effect of all the local demographic events across host populations are taken into account. In our study system Plantago lanceolata is a host to the powdery mildew, Podosphaera plantaginis, an obligate wind-dispersed parasitic fungus. These studies are conducted in the Åland Islands where the distribution of P. lanceolata is highly fragmented and its spatial population structure has been closely recorded for the past decade. Pathogen dynamics are studied regionally as colonization and extinction events, locally in habitat patch networks, as well as within local host populations to obtain data to parameterize a model of pathogen occurrence and prevalence. These data are complemented with laboratory studies to understand how spatial dynamics are driven by differences in resistance-pathogenicity. Specifically, these studies are designed to test whether there is variation in the resistance structure of the host both within and among populations and how this structure links to the observed disease dynamics. Also, studies are conducted to test whether the pathogen is locally adapted and at what spatial scale. Results of these studies provide new insights into host-pathogen interactions at multiple spatial and temporal scales. Interactions between host plants and their fungi do not occur in isolation but are likely to affect other species simultaneously exploiting the same host. Previous work on this system has demonstrated negative effects of the powdery mildew infection on the larvae of Melitaea cinxia. Future studies will determine whether these negative effects reach the next trophic level, affecting the parasitoids of M. cinxia.
Key publications:
Laine, A-L. A powdery mildew infection on a shared host plant affects the dynamcis of the Glanville fritillary butterfly populations. Submitted manuscript. 24 - Projects Biology of an old adaptive radiation: Evolutionary community ecology of the beetle tribes Canthonini and Helictopleurini in Madagascar
Researcher: Ilkka Hanski and Gavin Hinten Undergraduate students: Helena Koivulehto and Heidi Viljanen
The study of adaptive radiations has a long pedigree in evolutionary biology, starting from Charles Darwin’s chance discovery of the Galapagos finches during his visit to Galapagos Islands in 1835. Dung beetles in the tribes Canthonini and Helictopleurini in Madagascar have great potential for studies of adaptive radiation, because they form a large group of strongly interacting (competing) species, and because their ecology and biogeography on the small continent of Madagascar can be worked out with a reasonable research effort. Dung beetles possess a distinct advantage for field studies as their quantitative sampling is easy and quick and can be done even by untrained field assistants. This new project was started in 2002. The specific objectives are (i) to construct a molecular phylogeny of Canthonini and Helictopleurini to investigate their evolutionary history, (ii) to record their current geographic ranges in Madagascar with systematic sampling, and (iii) to study the community ecology of the beetles, which is known to be greatly affected by strong interspecific interactions. Ultimately, all these components will be brought together, to analyse the dynamics of speciation, range shifts, and ecological adaptation in this large but ecologically well-defined group of species. This project will also contribute to questions about the biological consequences of loss and fragmentation of tropical forests, which are severe threats to biodiversity in Madagascar. Two undergraduate projects were started in 2003, on edge effect, habitat selection and diet choice of beetles. A Madagascar-wide sampling of beetles was initiated, with the aim of obtaining a sample from about 50 forest reserves across Madagascar. Sequencing of the species available so far has been started in 2003. The first results support the hypothesis of two unrelated clades in the tribe Canthonini in Madagscar.
Collaborators:
Prof. Patricia Wright, Stony Brook, USA Mr. Rahagalala Pierre, Antananarivo, Madagascar Mr. Olivier Montreuil, Natural History Museum in Paris, France Dr. Jason Mate, Natural History Museum in London, UK Projects - 25 Influence of climate change on the distribution and abundance of species
Researcher: Ilkka Hanski Post graduate student: Varpu Mitikka
Research conducted during the past 10 years has produced convincing evidence of the on-going climate change having a profound influence on the demography and distribution of species. It has been suggested that climate change induced distributional changes pose a threat to the persistence of species that is comparable to the threat caused by habitat loss and fragmentation. Research on butterflies in Europe has demonstrated how climate change and habitat change interact. For instance, it is more difficult for specialist than generalist species to move to new areas with changing climatic conditions because the habitat for the specialists is often highly fragmented. Those butterfly species that have expanded their northern range boundary in recent decades tend to exhibit evolutionary changes at the expanding front, including increased mobility in comparison with older populations well within the range limits. This is a new project started in 2003. We have selected the map butterfly (Araschnia levana) for empirical studies. This species was first observed in Finland in 1973, and has started to spread from two directions, east and south, since 1983 (see p. 44). The purpose of the research is to model the range expansion and to study possible evolutionary changes that may have occurred during the expansion.
The map butterfly, Araschnia levana. Photo by Tari Haahtela. 26 - Projects
Biodiversity in boreal forests
Researchers: Ilkka Hanski, Otso Ovaskainen, and Gergely Várkonyi Post graduate students: Jenni Hottola, Laura Kivistö, and Reijo Penttilä
The year 2002 was the final year in our boreal forest biodiversity project funded by the Finnish Biodiversity Research Programm (FIBRE, 1997-2002), though research on forest biodiversity will continue at a smaller scale also in the coming years. Doctoral dissertations have been/are being completed on the biology of Xestia moths (Várkonyi; examination in September 2003), on the biology of bracket fungi (Penttilä; examination in the spring 2004, and Hottola; started in 2002), and on the biology of old-growth forest lichens (Kivistö; examination in 2004). Here we highlight three areas where much research has been conducted in the past years in this project. First, we have examined theoretically and empirically the extinction threshold phenomenon both at the stand level and at the level of larger forest areas. For instance, our studies have conclusively shown that the extinction threshold for threatened saproxylic fungi is around or beyond 20 m3 of coarse woody debris ha-1. Second, we have studied both theoretically and empirically the time delay in the response of species to changing forest structure. Modelling studies show that the time delay is longest in the case of species close to their extinction threshold (that is, species that are currently threatened), which is likely to lead to an underestimate of the extinction debt at the community level. Third, we have examined the genetic variability of old-growth forest populations and species: Xestia moths, the endangered beetle Pytho kolwensis and two of its relatives, the flying squirrel, and the trembling aspen.
Key publications:
Hanski, I. and Ovaskainen, O. 2002. Extinction debt at extinction threshold. Conservation Biology 16, 666-673.
Pakkala, T., Tomppo, E. and Hanski, I. 2002. Spatial ecology of the three-toed woodpecker in managed forest landscapes. Silva Fennica 36, 279-288.
Collaborators:
Mr. Juha Siitonen, Finnish Forest Research Institute, Helsinki Prof. Erkki Tomppo, Finnish Forest Research Institure, Helsinki Mr. Timo Pakkala, Natural History Museum in Helsinki Personnel - 27 MRG personnel and their research interests Mar Cabeza, Post graduate student
What is the minimum combination of areas needed to protect at least one (or a few) population of every species? – This question exemplifies the problem of selecting a system of reserves. Computational methods to solve similar questions have been developed during the past 30 years, however most of the approaches have focused only on representation of biodiversity, ignoring how well biodiversity persists once a reserve network is established.
In my recently completed thesis I have shown how poorly can simple reserve design methods perform in the long term if species spatiotemporal population dynamics are ignored. Following this we have developed different approaches that deal with spatial dynamics and habitat loss in various ways (see thesis section). The figure below shows that, when spatial population dynamics are taken into account, the optimal selection of reserves (number and spatial configuration) may also vary according to different assumptions of future habitat loss (from no alteration in (A) to complete loss of unselected sites in (C)).
Example for 19 butterflies at the Creuddyn Peninsula (UK). Selected sites shown in black.
The next step requires moving from extreme assumptions of habitat loss to more realistic modelling of future changes in the landscape. I am currently addressing the question of how to incorporate effects of changing landscapes into reserve selection methods. Such question involves assessing different problems, e.g. natural changes such as succession; habitat transformation (loss) at the non-selected areas; habitat changes due to climate change, etc. 28 - Personnel Oscar Gaggiotti, Senior researcher
Understanding the patterns of genetic variation observed in natural populations requires a mechanistic integration from the molecular biology of individuals through the genetics and ecology of whole populations and the evolutionary analysis of historical contingencies that these mechanisms may entail through random drift.
The integration of ecology, demography, and genetics has to occur at both the theoretical and empirical level. Theoretical integration is best achieved through the development of models that incorporate the interaction between demographic, ecological and genetic factors. The integration in the empirical area requires the development of sampling schemes that focus on both ecological and genetic attributes of populations, and the development of sophisticated statistical approaches that can utilise the information provided by different types of data (genetic, demographic, ecological, etc.) In line with this philosophy, my research covers a broad range of areas that include ecology, population genetics, evolution, conservation biology and statistics. Its main objective is to bridge the gap between population genetics and population ecology by extending existing evolutionary and population genetics theory and statistical methods to include more realistic ecological scenarios. The objective of this research is two-fold. Firstly, to understand the evolutionary processes that lead to the patterns of biodiversity observed in natural ecosystems. Secondly, to provide statistical tools that use genetic, ecological and demographic data to study metapopulation processes. The new insight thus acquired will help design management strategies that will maximise the amount of genetic variability maintained by populations that are affected by ever increasing human disturbances. Personnel - 29 Olivier Gilg, Post graduate student
My PhD work, started in 1998, investigates the lemming cycle and predator-prey interactions. Several recent studies have converged to the conclusion that the periodic rodent oscillations in boreal and arctic regions are essentially maintained by the interactions between the rodents and their mustelid predators. However, before arriving at a definite conclusion for the 70-yr- old puzzle of rodent dynamics, several hypotheses need to be tested, combining appropriate empirical and theoretical approaches. For this purpose, I’m adapting and testing the empirically-based models developed by Hanski and co-workers during the past ten years on a simple and well-documented high arctic vertebrate community from northeast Greenland. An ongoing collaborative effort between the universities of Helsinki, Montpellier and Freiburg was initiated in 1998 in order to collect information needed to parameterize the model.
Having documented the numerical and functional responses of the main lemming predators (stoat, Arctic fox, snowy owl and long-tailed skua), we are now able to estimate the summer predation rate for the entire predator guild, according to the lemming density at snowmelt (N’) and to the current lemming density (N). This predation rate is delayed, in line with the specialist predator hypothesis. Before the lemming peak (left panel) the predation rate is only high if lemmings are abundant at snowmelt (gray surface: 1% threshold at which predation rate equals lemming recruitment). After the peak phase (right panel), the predation rate is higher due to delayed density dependent predation of the stoat (that maintains the long-lasting low phase of the lemming cycle by preventing the prey population to recover). These functional and numerical responses were used to parameterize a predator-prey model that predicts a cyclic dynamic of similar amplitudes and cycle lengths as observed in the field.
All species before All species after
15 the peak phase 15 the peak phase 14 14
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e 11 11
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i
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y
y 4 4
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i 3 3 Da Da 2 1 2 1 1 1 10 N 10 N 0,1 0,1 1 1
0,01 0,01 N' 0,1 N' 0,1 30 - Personnel
Sofia Gripenberg, Post graduate student
I started my PhD studies in the MRG in June 2003. My project focuses on the interactions between the leaf-mining moth Tischeria ekebladella and its host plant, the English oak Quercus robur. I will examine in what ways variation in leaf quality and the distribution of oak trees in the landscape affect central processes in the population biology and evolution of the moth. The work is an extension of my MSc thesis (completed in 2002) that dealt with the spatial population structure of T. ekebladella and the adaptation of local populations to their host tree individuals.
In the summer I visited all oak trees on the island of Wattkast in south-western Finland. A database with information about the approximately 1600 oak individuals on the island was built and will form a base for future surveys and experiments in my PhD project. In September I re-visited a selection of the trees to investigate the distribution and abundance of T. ekebladella. As expected, mines were less likely to be present on isolated trees. In addition to this survey, I did a more extensive study of twenty oak trees growing in fairly close proximity in an area of high tree density. The greatest variation in mine density was found among leaves and shoots within individual trees, suggesting that female moths may be able to track variation in leaf quality at very fine spatial scales.
The distribution of oak trees and T. ekebladella on Wattkast in September 2003. Black dots depict trees that were not surveyed, white circles surveyed trees from which T. ekebladella was absent, and grey circles surveyed trees on which T. ekebladella was present. Mines were found on 73 of 113 investigated trees. Personnel - 31 Tapio Gustafsson, Research secretary
I am working as a research secretary for Professor Ilkka Hanski. Part of my work is to take care of the computing hardware and software used in the MRG. I also produce illustrations for different purposes and it is on my responsibility to update our website. I help to co-ordinate our group activities and write reports etc.
Last year our move to Viikki was the big thing and many of our computers were updated to Win2000 operating system. This year we got our first computers with WinXP, again a new thing to learn. At this point I can say that I have learned the basics of it. Laptops have been very popular and many of us have bought one for themselves. Another technical move is the gradual change to flat screens. Both things make users life easier and more flexible but laptops are a bit more complicate to manage compared to the desktop computers. At the end of the year I had a great oppurtunity to take part to the Madagascar Expedition. I learned a lot about the dung beetles and the work in the tropical rain forest. Unforgettable experience.
Madagascar is famous of its endemic chameleons. Photo by Tapio Gustafsson. 32 - Personnel Sari Haikola, Post graduate student
Inbreeding is expected to be frequent in small local populations of Melitaea cinxia in the Åland Islands. Therefore, selection during continuous inbreeding could in theory have purged the genetic load due to deleterious recessive alleles from the metapopulation. Our laboratory study shows that even though purging has to some extent occurred, part of the genetic load still remains in the system. In the laboratory, several different traits at various life stages show strong inbreeding depression. These traits include important fitness components such as egg hatching rate and larval survival. In brief, inbreeding depression is manifested over the entire life-cycle of M. cinxia. As there is no discrimination against close relatives as mates, inbreeding depression is expected to be an influential factor also in natural populations.
Saccheri et al. (1998) found that increased homozygosity was associated with elevated extinction risk of small populations of M. cinxia. In our laboratory study butterflies originating from old small populations in Åland were on average more homozygous than butterflies from small newly-founded populations. Similarly, parental relatedness was highest in the old small populations, which was associated with lowered egg hatching rate, larval survival and overall fitness in this population class as there was a negative relationship between parental relatedness and offspring fitness. Our laboratory study has clarified the relationship between homozygosity and extinction risk of small natural populations. The overall conclusion is that inbreeding is likely to have a substantial impact on the dynamics of the M. cinxia metapopulation in the Åland Islands. Personnel - 33 Ilkka Hanski, Research Professor and Director of the MRG
My research interests include population and community ecology and conservation biology, but for the past 15 years the real excitement has been, and still continues to be, metapopulation biology. I also have a commitment to make a contribution to conservation politics, in the form of popularizing research results, especially in the context of forest conservation in Finland. During the past year, I spent much time in completing two edited volumes, one on metapopulation biology (edited with Oscar Gaggiotti) and the other one on the biology of checkerspot butterflies (edited with Paul Ehrlich).
In 2003, I initiated a project on tropical forest dung beetles in Madagascar. I did my PhD on British dung beetles in the late 1970s, and I had a chance to work on tropical forest dung beetles in Sarawak (Malaysia) in 1978 and in Sulawesi (Indonesia) in 1985. The focus of that research was in community ecology, mostly in terms of spatial patterns, niche relationships, and abundance distributions. The community of dung beetles in tropical forests is extremely competitive: most resource, dung and carrion, is consumed in a matter of hours, and I was curious to see what sort of consequences the intense competition would have on community structure. In the new studies in Madagascar, I plan to return to some of the same questions, but with two new sources of information, a molecular phylogeny for the species (which has to be prepared) and knowledge of the geographic ranges of the species (based on large-scale sampling that has already been started). The Madagascan dung beetles comprise two large endemic radiations with around 300 species. I consider that there is much potential in merging phylogenetic and community ecology studies in the context of geographical ecology. The years to come will show what we manage to get done with the example of forest dung beetles in Madagascar.
Photo by Tomas Roslin. 34 - Personnel Paavo Hellstedt, Post graduate student
My Ph.D. research interests involved four aspects of stoat and weasel population biology: 1) Estimating the survival rate of captive-born least weasels in nature. We suggest that there are two main components explaining the differences of survival rate; the age of the animal, and time of year of release. 2) We also estimated the indirect effects of least weasel presence on Microtus-vole behaviour and demography. 3) In Lapland, we studied causes of change of the microtine cycles and the dynamics, habitat and prey selection of stoats and weasel. We suggested that change in mustelid densities refleced change in microtine populations. 4) Also we started (with Pekka Helle from the Finnish Game and Fisheries Research Institute) a new project about large scale spatial and temporal dynamics of the stoat and weasel populations.
Using radio-tracking, we estimated the behaviour and the survival of the captive- born least weasels (Mustela nivalis nivalis) released in nature. We equipped with a radio-transmitter 27 captive born weasels and six caught on site, marked and released weasels. We studied the survival of the different aged captive born weasels and differences in the survival in different seasons. Generally the daily mortality risk was higher with captive born weasels than with wild caught weasels. The survival rate of captive born weasels was higher when they were released during the summer and hence they were younger than during the other time of the year. Also, we studied home ranges of the weasels, weasel’s use of terrain, visibility and avoidance of predation. Differences in the behavioural pattern of captive born weasels and wild caught weasels were apparent. Captive born weasels were more often visible on the top of the ground vegetation and were more undaunted than wild caught weasels. Personnel - 35 Gavin Hinten, Post doc
My research in Australia focused on a model system of bush rats (Rattus fuscipes) isolated on islands through rising sea levels since the last ice age, an ideal system for studying the genetic consequences of isolation and fragmentation. We were able to focus on populations where we know the size and time of isolation of the islands and where results are not confounded by gene flow. Further by using neutral microsatellite markers we were able to control for selection. Our results show that isolation and fragmentation have a double impact on genetic diversity. First, the small island populations have undergone extreme bottlenecks in the past, resulting in loss of genetic diversity, increased inbreeding, and presumably increased genetic load. Second, genetic drift has resulted in rapid genetic differentiation between populations. In fact, genetic drift is such a dominant evolutionary force in the small island populations that in a similar study of genetic diversity in a gene region of the Mhc, a region under strong selection, genetic drift has overridden selection in these populations.
I joined the MRG in 2002 to continue research into the genetic consequences of isolation and fragmentation, this time with metapopulations of shrews on islands in lakes. Shrews are well suited to studies of metapopulation dynamics. They have a high energy demand and metabolic rate yet low energy reserves, placing enormous pressure on colonisation and extinction processes. My research will focus on three shrew species, Sorex araneus, S. caecutiens, and S. minutus. These species overlap extensively in habitat but have distinct physiological, behavioural, and ecological adaptations that allow them to survive in a temporally changing environment. In particular there is little overlap in body size among these species. However body size plays an important role in shrew metapopulation dynamics. Larger bodied shrews have greater dispersal ability, greater resistance to environmental stochasticity, and are able to dominate inter specific competition for habitat patches. Alternatively smaller bodied shrews are better colonisers and are able to survive in less productive habitats, possibly because they have a lower per capita food requirement. My aim is to integrate patterns of genetic diversity with available ecological and physiological data from island and mainland metapopulations of these shrew species to gain a better understanding of the genetic component of metapopulation dynamics and the evolutionary processes of these species. 36 - Personnel Jenni Hottola, Post graduate student
My main research interests are in forest ecology. I am especially intrigued by wood-rotting fungi, which are key species in many respects. They are particularly susceptible to alterations in forests due to their dependence on coarse woody debris and often specific requirements pertaining to microclimatic conditions and quality of the growing substrate. Currently, 37 % of polypores in Finland are classified as threatened or near threatened, mainly because of forestry practises.
My MSc thesis dealt with relationships between characteristics of coarse woody debris and communities of wood-rotting fungi, mainly polypores. I work as a researcher in the Finnish Forest Research Institute, where my task is to assess the state of polypores in forests of southern Finland and to examine the importance of new practises in forest management. The main objective of my PhD project is to examine the patterns and processes that characterise the dynamics and persistence of wood-rotting fungi on different spatial and temporal scales. On the scale of forest patches, the aim is to study how the structure of the forest patch affects the dynamics of local populations. In addition, species interactions will be studied to find out the quality and extent of their influence on community structure. On the regional landscape scale, I will study the spatial incidence of polypores in relation to the structure of the forest landscape and the processes operating on the landscape level, while also taking the long-term dynamics into consideration.
The figure shows one result from my MSc thesis, based on a partial canonical correspondence analysis of species, plots, and variables describing coarse woody debris. Each vector depicts the direction and strength of maximal linear correlation between the explanatory variable and the plot values. The result indicates that volume, diversity, and mean decay stage of logs have the biggest though dissimilar effects on the incidence of polypore species. Number of logs and standard deviation of decay stages of logs are of less importance. Personnel - 37 Maaria Kankare, Post graduate student
My PhD project includes studying the phylogenetic relationships among Cotesia (Braconidae) species parasitizing Melitaeini butterflies. Six Cotesia species from 16 host butterfly species from Europe, Asia and North America were examined using DNA sequence data as well as 12 microsatellite loci. Analyses using both maximum parsimony and maximum likelihood revealed two distinct Cotesia clades (A, B). In one clade (A; C. acuminata; C. bignellii) host ranges are apparently narrow, while in the second clade (B; C. melitaearum; C. lycophron; C. cynthiae) host utilization across the Melitaeini is more extensive and reflect a more recent, or active, speciation processes. Neighbour-joining trees calculated separately for the two main clades based on microsatellite allele frequencies were consistent with phylogenetic trees obtained from the sequence data. These results strongly suggest the presence of several additional, previously unrecognized, Cotesia species.
In the second study, the parasitoid assemblages associated with rich Melitaeini butterfly communities, focusing on the specialist Cotesia parasitoids, were characterized. Samples were collected from several sites in Catalonia, northern Spain, a region that harbours one of the richest butterfly fauna in the Europe. As a first step, intensive sampling in a few sites provided basic information on the natural history of most of the host butterfly species, including reliable data on hostplants and phenology, as well as on species co-occurrence, abundance and specialist parasitoids. Secondly, behavioral observations of female parasitoid host-acceptance in the laboratory were used to study the potential for host specificity and the possible presence of cryptic species among the Cotesia. Additionally, the host specificity and geneflow among Cotesia parasitoids were studied using microsatellite markers. Molecular and behavioral data will be used in combination with the natural history observations to investigate how closely linked ecologically the Melitaeini butterfly community may be.
Melitaea deione larva feeding on Plantago lanceolata. Photo by Constantí Stefanescu. 38 - Personnel Jonna Katajisto, Post graduate student
Some years ago I ended up in the Scandinavian Brown Bear Research Project (SBBRP) as an undergraduate student. After finishing my MSc thesis about habitat selection of female brown bears I joined MRG as a PhD student. Bear habitat selection is still one of my interests, but at larger spatial scale together with other spatial processes affecting the distribution of bears. I also continue in close collaboration with the Scandinavian bear project that provides the data.
My aim is to investigate the effects of landscape structure and human activity on brown bear dynamics and distribution in Fennoscandia by means of spatially explicit population models. In these SEPMs biological information of the species is combined with real-world spatial information represented by maps of habitat quality and human activity. The work is based on exceptionally large long-term radiolocation data set collected by SBBRP. This material provides good information about demographic processes and habitat selection of bears. It also contains sequences of consecutive locations, which make it possible to study both detailed movements and lifetime dispersal. Results will be applied to Finland in collaboration with Finnish Game and Fisheries Research Institute. Essential processes to include in SEPMs are dispersal and home range establishment that link bears with their environment. Our results from habitat selection show that bears avoid human activity. Consequently, bear movements and home range (HR) establishment are more limited by human influence than by natural factors related to habitat quality. Using traditional HR methods in habitat selection analysis may result in loss of information or bias towards frequently observed areas when applied to data with irregular sampling intervals (e.g. data that includes intensive observation periods). We developed a new kernel-based HR analysis method, which includes kernel smoothing also in the time dimension.
Home ranges produced with traditional and new home range methods. (A) In kernel home range with resampling to avoid autocorrelation many observations are left outside the range and information is lost. (B) In time kernel home range no data is lost and some areas with high use outside traditional kernel are shown. Personnel - 39 Laura Kivistö, Post graduate student
Ever since I attended the basic lichen course in 1993, I have been interested in the magical world of lichens. Lichens are actually kind of mini-ecosystems consisting of at least two organisms: a fungus and a photosynthetic partner. My research focuses on the effect of old-growth forest fragmentation on epiphytic lichens. Actions to maximize tree productivity in managed forests, e.g. short rotation cycles, tree monocultures and even-aged tree stands, are the most important factors at stand level that affect epiphyte communities. Forest owners need effective tools to estimate the effects of forest management on biodiversity. Indicator species have been widely used to this purpose. Using old-growth forest lichen species I have examined 1) if isolation from a large old-growth forests affects the epiphytic lichens, 2) how different lichen species respond to old-growth forest fragmentation, i.e. are old-growth forest specialists more sensitive than common generalist species, and 3) what structural elements in the forest fragments are necessary for the survival of old-growth specialist lichens. The age of the dominant trees of the forest stand appears to be the most important factor affecting old-growth specialist lichens. The importance of stand age can not be emphasized too much; rotations should be prolonged in managed forests and more old-growth forests should be left unlogged to maintain the diversity of old-growth specialists. At stand level, some old and large tree individuals should be left in managed forests as “a legacy” for the next generations of several organisms. Especially old aspens and willows host a very rich community of epiphytes.
Lobaria pulmonaria is one of the old-growth forest indicator species in Finland. It grows usually on old Populus tremula and Salix caprea trees. Photo by Kimmo Jääskeläinen. 40 - Personnel Mervi Koskela, Post graduate student
In my PhD project, started in 2003, I concentrate on questions about the factors that influence the incidence of saproxylic species, i.e. species depending on dead trees. Beetles (Coleoptera), polyporous fungi (Basidiomycetes) and lately also slime molds (Myxomycetes) are of my interest.
There are several factors that have an effect on whether a species occur in a certain habitat patch or not: the amount and quality of dead trees, spatiotemporal continuity of suitable host trees, habitat loss, isolation connectivity etc. The spatial scale at which a certain process function is specific to species depending on dispersal ability and stage of specialization. I investigate the following questions: How does the amount of host trees affect the occurence of certain generalist and specialist species? Does the spatiotemporal history of the area have an effect? At which scale does spatial factors work? Entomology is an old discipline of science. Compared to other insect groups many species of Coleoptera are well studied and quite easy to find as their habitat requirements are known. This forms the basis for using direct search method when studying threatened beetles, which are usually rare in trapping samples. Some of the species are even quite easy to identify in the field, thus there is no necessity to collect specimens. During this year I have also researched slime molds and their associates, especially Agathidium pulchellum (Leiodidae). It has been thought that this endangered beetle species depends on dead aspens, but new data shows that actually aspen may not be that important for the species. My aim is to compare this species to the common species of the genus and resolve their life-cycles and interactions with Myxomycetes.
Larvaes of Agathidium pulchellum live on slime molds (Myxomycetes) of genus Trichia decipiens (in picture) on dead lying spruces. Photo Mervi Koskela. Personnel - 41 Anna-Liisa Laine, Post graduate student
The Plantago lanceolata meadow network in the Åland Islands provides the framework for my PhD studies, which are focused on an obligate pathogenic fungus (Podosphaera plantaginis) infecting a fraction of local Plantago populations. The host- pathogen dynamics in this system are strongly driven by the spatial configuration of the host; the local populations are abundant, generally quite small and fragmented in distribution. While the importance and diversity of plant pathogenic fungi have been recognized in many ecosystems, the dynamics of natural fungal populations remain poorly understood. In particular, to date almost nothing is known of why particular host populations are not infected and how the resistance structure of local host populations links to variation in the incidence and severity of natural disease epidemics.
This year, using an experimental approach, I studied a subset of the local Plantago populations to test whether adjacent host populations showed differences in the identity and frequency of particular host resistance phenotypes, and whether these differences are related to their disease status and spatial location. Results showed striking differences in the resistance structure both within and among host populations. Out of the 64 host individuals altogether 16 resistance phenotypes were identified. The frequency of the resistance phenotypes within the host populations was uneven - in the extreme case all studied individuals within a population represented a unique resistance phenotype. In the other extreme, 50% of the hosts within a population were identical in their resistance responses. Non- infected populations showed markedly higher mean resistance than infected populations, which may retard the spread of the fungus and thereby decrease the probability of regional epidemics.
(A) Resistance of the host plants within each of the eight host populations to the four P. plantaginis strains. (B) The mean level of resistance of the eight P. lanceolata populations as scored in the inoculation experiment. Infected populations are represented by dark bars 42 - Personnel Evgeniy Meyke, Research assistant
My primary task in the MRG is maintaining the database part of Cinxia survey and developing Database and GIS applications for the members of the group. Quite often the software for a specific task is not available or does not do exactly what people need. Developing such customized, standalone Windows applications, which combine database access and GIS mapping capabilities turned out to be very useful especially when dealing with problems posed by data coming from Cinxia survey. The example screenshot demonstrates one of the applications that manipulates and analyses data from MS Access database and ArcView’s shape files. Modern software development tools (such as Borland Delphi and C++ Builder) allow rapid application development with easy addition of many useful features (e. g. powerful reports, visualisations, graphs, etc.). Field forms for Cinxia survey are automatically printed from specially built GIS/Database application and include detailed information facilitating the work of surveyors. Subsequent automatic scanning of field forms directly into the database has allowed for broader range of data to be recorded and has replaced manual entering of data via keyboard. I am also in the process of completing my Master’s thesis in the Metapopulation Research Group, based on collection of Parnassius species from Kyrgyzstan in July 2002 and 2003. Personnel - 43
Atte Moilanen, Senior researcher
Currently I pursue three lines of research. Foremost of these is the rapidly expanding reserve network design project, which is about the application / incorporation of metapopulation dynamics to reserve selection (see projects, and pages for Cabeza and van Teeffelen). Personally, I am most interested in the development of new analyses, optimization methods and computational tools for reserve selection purposes. My second line of research is supervison of / collaboration with Jonna Katajisto in the modeling of the dispersal and spatial dynamics of the brown bear (see Katajisto´s page). I also continue work concerning metapopulation models and their parameter estimation (in collaboration with prof. Ilkka Hanski, Dr Otso Ovaskainen, prof. Juha Alho and prof. Chris Thomas´ group). The picture below (right) shows a recent result about the convergence of optimization in a nonlinear reserve selection problem, in which landscape connectivity (which depends on the spatial pattern of the selected reserve network), affects species- specific probabilities of occurrence in grid cells. This problem has a huge search space and is quite difficult for a simple stochastic search method based on simulated annealing or genetic algorithms (upper 10 replicates). The method that I have developed combines the stochastic global search (GA) with local search (LS) and a self-adaption of the control parameters of the GA and LS. This method converges quickly and consistently (lower 10 replicates) to the optimal reserve configuration, and the intention is that derivatives of this method will be used for handling the increasingly complex optimization tasks that we are encountering in reserve selection optimization. 44 - Personnel Varpu Mitikka, Post graduate student
I joined the project “Influence of climate change on the distribution and abundance of species” in October 2003. Many recent studies have reported on likely range shifts of species in response to changes in climatic conditions as well as changes in the availability of suitable habitat. Several insect species showing a northward range shift apparently in response to increasing temperature have been studied in Europe and North America.
The species of interest in my study is the map butterfly (Araschnia levana), which has expanded in Finland during past decades. The first observation of the species was made in 1973, and the first local population was found in eastern Finland in 1983 (see the map). In southern Finland, the map butterfly was first recorded in the beginning of 1990 ‘s, and it has expanded throughout the southern coastal areas in the past five years. In the first project of my study I will explore the changing distribution of the map butterfly in Finland using a modelling framework developed for similar research in the UK. In the model, distributional data for the species, data describing habitat suitability as well as selected climatic variables will be integrated to generate predictions for the distribution of the species under given circumstances. Detailed studies of the biology of the species are needed to understand the influence of any additional factors on the distribution and changes in it. Field studies and experiments will be conducted in summer 2004 to investigate habitat selection and spatial population structure. I will compare populations located in the expanding front versus in areas where the species has occurred for a long time.
The distribution of the map butterfly (Araschnia levana) in Finland 1973-2003. The site of the first observation (1973) is located between the two distinct distribution areas. Personnel - 45 Marko Nieminen, Senior researcher
My main responsibility within MRG is to keep running the monitoring and other field studies on the Glanville fritillary butterfly Melitaea cinxia in the Åland Islands, SW Finland. The main aims of this research are, firstly, to study different causes for the spatially correlated dynamics in M. cinxia metapopulations, and the spatial scales at which they operate. Two factors are apparently driving these spatially correlated dynamics: weather (fast changes with varying extent) and changing landscape structure (slow changes with relatively stable extent). They also correlate with several other factors, e.g. interspecific interactions, inbreeding effects and habitat quality. Secondly, to monitor all known suitable habitat patches bi annually. We collect data on numerous habitat characteristics and larval nests every autumn, and on the numbers of larvae and parasitoids in the spring.
Increases and decreases in the numbers of occupied habitat patches within subareas of Åland from 2002 to 2003. 46 - Personnel Saskya van Nouhuys, Senior researcher
The focus of my research is the link between large and small- scale ecological processes for interacting species. Central to this work is the roles of dispersal and habitat distribution in population dynamics and community structure. As part of the MRG (since 1997) I work primarily on the parasitoids associated with the Glanville fritillary butterfly in Åland, but over the past few years my interests have spilled out into the host-parasitoid relationships in Melitaeine butterfly communities throughout their range.
In 2003 I finished a long term study showing that the developmental synchrony between the butterfly Melitaea cinxia and the parasitoid Cotesia melitaearum varies spatially and temporally, depending on temperature. This variation has metapopulation-wide consequences for the parasitoid in Åland (see figure below). An ongoing project that I enjoyed working on in 2003 is the relationship among eight Melitaeine butterfly species and their parasitoids in Catalonia, Spain. Initially we thought that the butterflies would be tightly linked by shared Cotesia parasitoids, but field collection, adult parasitoid behavioral observation, and microsellite markers indicate that instead, the Cotesia are for the most part host specific. Interestingly the abundances of the ecologically similar co-occurring butterflies appear to vary independently, again suggesting that they are not linked by shared parasitoids. The single exception is the only pair of species that do in fact share a Cotesia parasitoid. I plan to look further into this during the next year and rate of colonization of host populations by the parasitoid Cotesia melitaearum (from van Nouhuys & Lei, JAE, in press).
The relationship between early spring temperature and parasitoid colonization rate. Personnel - 47 Toshka Nyman, Laboratory technician
My task as a laboratory technician is to facilitate the laboratory work of the Molecular Ecology Team by providing the needed materials, chemicals and enzymes. Extracting, amplifying and sequencing DNA is the biggest part of my every day job. This year we were happy to get a second sequencing machine and two gradient machines, which means less time for optimizing the conditions of PCR and practically no more queues for a sequencing machine.
Female Melitaea cinxia. Photo by Tapio Gustafsson.
Melitaea cinxia feeds the nectar of Allium schoenoprasum in June. Photo by Tapio Gustafsson. 48 - Personnel Otso Ovaskainen, Senior researcher
I completed my PhD in mathematics in 1998, after which I have been doing research in mathematical biology in the MRG. My main emphasis has been in the development of such metapopulation theory that can be used to assess the dynamics and persistence of assemblages of local populations inhabiting fragmented landscapes. During this year, I attempted to merge metapopulation level processes with individual level processes. To do this, I developed with Ilkka Hanski individual-based models (IBMs), which stemmed from our work in dispersal theory. We assumed that the dispersal phase may be characterized using correlated random walks and edge-mediated behaviour, which we have been studying with Stephen Cornell from Cambridge, UK. By supplementing the dispersal models with models of local population dynamics, we obtained a unified modelling framework for patchy populations with much movement among habitat patches and classic metapopulations with infrequent movements. I then derived stochastic patch occupancy (SPOM) approximations for the IBMs with the aim of building links between individual-level parameters and metapopulation level-parameters. The SPOMs derived from IBMs have parameters that related directly to the life-history and behaviour of individuals, which is an advantage for model interpretation and parameter estimation. Unlike the original IBMs, the SPOMs are analytically tractable and relatively parameter-sparse, which is an advantage for the development of general mathematical theory and for parameter estimation.
As shown by the figure, the SPOM approximations (dots) mimic well the behaviour of the IBMs (lines) when applied to a real heterogeneous network of 51 habitat patches inhabited by a Glanville fritillary butterfly metapopulation. Panel A assumes pure demographic stochasticity, panel B uncorrelated environmental stochasticity, and panel C spatially correlated environmental stochasticity. Personnel - 49 Reijo Penttilä, Post graduate student
My PhD project deals with the effects of forestry and forest fires on wood-rotting fungi, polypores in particular, and my studies focus on both the community structure and on the population structure and population dynamics of old-growth forest fungal species. I have also examined the dispersal ability and genetic structure of fungal populations. My studies have been part of the project “Biodiversity in Boreal Forests” and during the years 1997-2001 I worked as a senior researcher at the Research Station of Friendship Park in Kuhmo, where I was also organizing several other studies belonging to the project. At the moment I work at the Finnish Environment Institute, where my main task is to study, how restoration of boreal forests affects species dependent on dead wood.
In a recent study (Penttilä et al. 2003), in which I 8 compared polypore communities in old, managed and 7 Managed 6 Oldgrowth old-growth spruce forests in south-western Finland, 5 it was found that threatened polypore species were 4 almost totally confined to old-growth forests and were 3 2 only found from forest stands in which the amount of 1 3 0 dead wood exceeded 20 m /ha (see picture); this Number of threatened species figure appears to represent a stand-level threshold 1 10 20 100 200 3 value for the amount of dead wood, below which the Volume of dead wood, m /ha persistence of threatened species becomes unlikely. Extensive dispersal studies of Phlebia centrifuga, a threatened old-growth forest fungus specialized on large fallen spruce trunks, showed that spore dispersal is strongly concentrated to the close vicinity of fruitbodies, which probably prevents P. centrifuga from maintaining viable populations in highly fragmented landscapes. This conclusion is also supported by field inventories, which showed that the frequency and abundance of P. centrifuga is much lower in more fragmented old-growth forests of Pohjois-Häme (south-western Finland) than in less fragmented old-growth forests of Kuhmo (eastern Finland). A similar pattern was also observed for several other old-growth forest species (e.g. Amylocystis lapponica, Fomitopsis rosea), which indicates that the viability of populations of several old-growth forest fungi is strongly threatened by the scarcity of suitable habitat in southern Finland.
Penttilä, R., Siitonen, J. and Kuusinen, M. 2003. Polypore diversity in mature managed and old-growth boreal Picea abies forests in southern Finland. – Biological Conservation, in press. 50 - Personnel Juha Pöyry, Post graduate student
The main objective of my PhD work is to empirically examine the effects of cattle grazing on insect populations in semi-natural grasslands. My target taxa are butterflies, macromoths and aculeate wasps. The main theoretical framework of my work stems from the >dynamic equilibrium model= (DEM) predicting diversity patterns along gradients of productivity and disturbances and from metapopulation theory. In the year 2003, I progressed with my thesis by determining and analysing materials collected in previous years and by compiling manuscripts.
Results on the effects of restarted grazing on community structure and abundance of individual species, respectively, were reported in two submitted manuscripts. Species richness and abundance of butterflies and moths were generally higher in non-grazed than in grazed grasslands. In contrast, relative community diversities for butterflies and moths were highest in old pastures. More individual species occurred most abundantly in abandoned pastures than in old pastures, but those species that concentrated in old pastures had more often declined in their occurrence in Finland. Furthermore, it appears that the studied taxa respond differently to restorative grazing on community level (Fig.). Vascular plants in grasslands clearly benefit from continued and restarted grazing, whereas many insects have problems under the current high grazing intensities. These observations are qualitatively in accordance with the predictions of DEM.
Species density of vascular plants and species richness of pollinators (aculeate wasps and hoverflies), micromoths, and butterflies and macromoths among the three pasture types with differing management histories. Personnel - 51 Tomas Roslin, Senior researcher
My main interest lies at the interface between evolutionary ecology and spatially oriented population biology. Working primarily on insects, I assess where in the landscape given species occur, how tightly linked the local populations are by dispersal, and how this affects the evolution of local adaptations. In my current research project, Spatial population dynamics and adaptation in a host-herbivore system, I am using leaf-mining and galling insects on English oak (Quercus robur) as a model system. From these host-specific insects’ point of view, individual oak crowns represent islands of suitable habitat in a sea of unsuitable habitat. Hence, I assess how variation in the configuration of the host-tree archipelago affects central population processes among the insects.
One of the most intriguing results this year has been the quantification of qualitative variation in oak foliage at different hierarchical levels. In the graph below, I have partitioned random variation in the larval performance of a leaf-mining moth, Tischeria ekebladella, into variation between trees, variation between bagged branch tips, and variation among leaves. The error bars show approximate 95% confidence limits. Of the total variation in moth performance, only an average of 5% occurred at the tree level, and there was much more variation among leaves than among either bags or trees. This finding has at least three important implications. First, the similarity in average resource quality among trees renders classical metapopulation models a useful tool in understanding the population dynamics of specialist herbivores across larger sets of host tree crowns. Second, little variation between trees combined with large variation within trees makes local adaptation at the level of host tree individuals unlikely. Instead, insects may evolve to select their resource at the level of single leaves within trees. Third, large variation within trees calls for extensive replication within trees in ecological sampling designs and ultimately, it even questions the biological relevance of resolving minor differences among trees as compared to major differences among different parts of one and the same tree. 52 - Personnel Marjo Saastamoinen, Research Secretary
I joined the MRG in the beginning of October 2002 after finishing my MSc degree. In 2003 I have worked in the group as a research secretary replacing Anu Väisänen who has been on maternity leave. My primary duties include taking care of financial transactions, the budget, administrative duties as well as helping prepare proposals and reports. During the year, one of the most demanding tasks was to prepare an EU-proposal for the sixth Framework Programme.
During the summer I also started my post-graduate studies, which I will continue full-time after Anu returns. The main focus of my thesis will be dispersal of the Glanville fritillary butterfly (Melitaea cinxia). Various factors are known to affect butterfly dispersal behaviour and recent studies have suggested that fragmented habitat could select for increased flight propensity. It has also been shown that butterflies in newly-established local populations are more dispersive than butterflies in older local populations of the Glanville fritillary. The main purpose of my study this summer was to test whether individuals from newly-colonised populations have increased flight abilities and if so, can we find any trade-offs between dispersal ability and reproductive success. For the experiment, larvae from 20 newly-established and 20 older habitat patches were collected around the Åland Islands. After emergence from the pupae the individual butterflies were marked and released into a large butterfly cage (30 m x 30 m) covered by a mesh, constructed for the experiment on a natural habitat patch. The cage was divided into 64 grids and the butterflies were monitored systematically during their active hours by recording their location. The experimental set-up also allowed us to observe matings and female ovipositions thus enabling us to measure the reproductive success of each female and male butterfly. We use genetic markers to identify the parents of the few egg groups where oviposition was not observed directly. The experiments carried out in 2003 demonstrated that this kind of experimental set-up works perfectly for M. cinxia and allows us to conduct a variety of similar experiments in the future.
Photo by Niclas Fritzén. Personnel - 53 Tarja Salmi, Post graduate student
In my PhD project, I will examine the population biology of aspen (Populus tremula) in both old growth and managed forests. I’ll work at three different levels. At the level of individual tree and stand, I’ll study the reproductive biology of aspen. At the landscape level, I’ll assess the extent of and the reasons for the spatial aggregation of mature aspen trees.
My work is based on spatially referenced data on ca 30,000 mature aspen trees. The data shows that there is 5.5 m3/ha aspen wood in old-growth forest compared to the 0.8 m3/ha in managed forest. In old- growth forest more than 70 % of aspen wood is alive and 25 % is dead. In managed forest the situation is vice versa. In fact, in this 4000 ha of managed forest one third of dead aspen wood was destroyed by man. Including some additional data, I now have information on all trees (diameter >1 cm at breast height) within 65 and 30 ha of old growth and managed forests, respectively. I’m studying the spatial aggregation and dynamics of coarse woody debris as well as the regeneration of aspen. I’m also studying the age distribution and history of stand development. Preliminary results show that aspen populations are declining in old growth forest. In summer 2001 we did (with Leena Suvanto, see p. 56) a field experiment on the emergence and survival of aspen seedlings under varying moisture conditions. Results show that watering and sowing time affected the number of seedlings. The survival was low (10 %) and it depended on watering. Also seedbed conditions affected the emergence and survival of the seedlings. Thus we conclude that successful sexual reproduction of aspen is possible in nature if the right seedbed conditions with enough moisture are met. I started my maternity leave in September 2002 and will continue my work after that. 54 - Personnel Alia Sarhan, Post graduate student
In the framework of my PhD studies, I am studying the population genetics of the Glanville fritillary (Melitaea cinxia) on the Åland Islands, where it occurs as a typical metapopulation. In this system, the frequent extinctions and small population sizes might lead to a loss of genetic diversity and ultimately result in a lack of adaptability to changing environmental conditions.
There are several factors that are crucial in determining the strength of genetic drift, and hence the rate at which genetic diversity is lost. Among them we have inbreeding, biased mating success, gene flow between subpopulations, and size and composition of colonization propagules. In order to study whether inbreeding levels found in natural populations influence the butterfly population dynamics, we conducted mating experiments in the laboratory, using butterflies from small, isolated populations. Butterflies were mated either within family or among populations. Fitness components, like egg- hatching rate and larval survival were measured. These offspring fitness measures were correlated with the expected offspring heterozygosity, based on the parental genotype at five highly variable microsatellite loci. We found a very strong correlation. Survival increased with increasing heterozygosity. Biased reproductive success is another important factor that influences the loss of genetic variation. I would like to measure it from the field experiment we did in the Åland Islands this summer. For that, I will first have to determine the parents of the larval groups for which mating and oviposition were not observed. Another important factor is the size and composition of colonization propagules. I have sampled newly colonized populations from areas of different butterfly densities, and I will try to estimate how many colonizers founded these populations and how related they are. 0.8
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0.10.30.50.7 exp. homozygosity Personnel - 55 Janne Sundell, Post doc
My research interest focuses on the role of predation on the population dynamics of small mammals. To obtain better understanding of the effects of predation, I have concentrated both the population dynamics of the prey and predator separately by describing the spatial and temporal patterns in their dynamics. These studies include the investigation of the large-scale spatial dynamics of voles as revealed by the breeding success of vole- eating avian predators. Currently, I am working on the spatial and temporal dynamics of small mustelids, the main predators of voles. My aim is also to investigate the connection of these dynamics to the local prey dynamics. Despite the use of these extensive data sets in my research, I consider my self as an experimentalist. I have used the experimental approach in most of my studies. These experiments are conducted in the laboratory, large outdoor enclosures and in natural environments. The experimental approach is used in studies on functional and numerical responses and the prey choice and hunting behaviour of the least weasel. With aid of the results of the laboratory experiments and observations in nature, together with K. Norrdahl (Univ. of Turku), we developed an alternative predation- based explanation for the Chitty-effect, the phenomenon that describes phase-related changes in the mean vole size. The main experiment of my thesis work, completed in 2002, was a large-scale field experiment testing the predation hypothesis. During my short postdoctoral visit in University of Jyväskylä in 2002, I focused on the antipredatory behaviour of the prey. I studied the refuge use of voles under the predator presence and how voles allocated their time between the foraging and vigilance under the temporally changing predation risk. Studies involved also the investigation of changes in the stress levels and feeding behaviour of voles when exposed to the predation. Recently, I have started a project, in which the behaviour of the brown bear will be studied with the method combining the modern GPS-satellite tracking and GSM- mobile phone techniques. My aim is to boldly 0.6 study how bears behave in human contacts. 0.4 Figure shows the relationship between the 0.2 degree of synchrony (mean ± SE cross- correlation coefficient with lag zero) and 0.0 distance in Finnish vole population of synchrony Degree fluctuations as revealed by the breeding -0.2 success of vole-eating avian predators. 0 200 400 600 800 1000 1200 Distance (km) 56 - Personnel Leena Suvanto, Post doc
The first three and a half months of year 2003 I worked as an assistant co-ordinator in EU proposal “Method for Environmental Risk Assessment of Alien Genes in Forestry”. This proposal was submitted to the VI Framework of European Union for the sub- priority “Global Change and Ecosystems” as an Integrated Project. It had 21 participants from 12 different countries and the total budget of the proposal was nearly 12 Meuro. Unfortunately the proposal was not financed, but the experience was educational and interesting.
The rest of the year I worked in a new project “Gene flow in aspen (Populus tremula)”, which is a co-operation project with Finnish Forest Research Institute (see page 22). In year 2003 the research was focused on studying the possibility of gene flow between hybrid and native aspen. My work was concentrated on analysing the genotypes of adult and juvenile trees using microsatellites. The origin of the seedlings could not quite be determined with microsatellites, which was mainly due to lost mother genotypes (e.g. logging). I am also using the same microsatellites to analyse the competition cross results between aspen and hybrid aspen. The work is still in progress, but we are trying to find out whether the species have different competing ability in pollen mixtures. Preliminary results show that the females could somehow choose which species’ pollen is used to fertilise the ovules. I also supervised the lab work of Hanna Alanen, who did her Master thesis in the Department of Forest Ecology on aspen clone identification using morphological, phenological and genetic markers. This work was done on assignment by the Plant Production Inspection Centre, and is connected to the new laws regulating the marketing of forest Mean germination % in different species reproductive material. combinations
70 60 50 40 hybrid mother % 30 tremula mother 20 Mean germination % 10 of seeds originating 0 from crosses between hybrid father father mixture tremula father aspen (P. tremula) and father hybrid aspen. Personnel - 57 Astrid van Teeffelen, Post graduate student
I joined the MRG as a graduate student from Wageningen University, the Netherlands, in 2002. I returned to the MRG in September 2003, as a post graduate student. Predicting the occurrence of species by means of so-called habitat models, which can be implemented in reserve selection procedures, is currently my main interest.
It is often difficult or costly to obtain occurrence data for multiple species at a scale at which reserve design can be applied. Even when occurrence data would be available, it is often collected in a single time period. Such data is a snapshot of species’ occurrence and does not provide information about where species are more likely to occur, which is crucial information for successful reserve design. Models, which predict the occurrence of species, can be useful tools in distinguishing between patches for their contribution to population persistence. Habitat models generally use information about habitat suitability to predict the species’ occurrence in a landscape. These models can be made more ecologically realistic by implementing measures of population dynamics, and connectivity measures. During this first part of my PhD I have surveyed literature concerning connectivity, as well as habitat modelling in general. With this knowledge I would like to implement connectivity measures in habitat modelling. Further, I want to investigate the different types of existing habitat models and find the most suitable models for implementation in reserve network design.
The figure shows two maps of a study area in the Netherlands: Panel A shows abundance data of the black woodpecker Dryocopus martius (white to black, 0 to 4 recorded breeding pairs). Panel B shows the predicted occurrence of black woodpecker, based on habitat suitability and two connectivity measures (white to black: probability of occurrence ranges from 0-0.2 to 0.8-1.0). 58 - Personnel Gergely Várkonyi, Post graduate student
My research interests lie in (i) population biology of insects with periodic dynamics, (ii) conservation biology of organisms dwelling in boreal forests and (iii) research on parasitic wasps, with special reference to the family Ichneumonidae.
In 2003 I completed my PhD thesis on Population biology of periodic Xestia moths (see Theses). From 1998 onwards I have been coordinating mutual conservation biology studies of the MRG and the Research Centre of Friendship Park in Kuhmo, eastern Finland (see Field sites). One of the major projects focused on changes in species richness on small old-growth forest (OGF) fragments after the isolation of the forest patches. Increasing levels of spatiotemporal isolation of the fragments affected OGF organisms in different ways, e.g. species richness of highly specialized lichens decreased gradually, and the frequency of OGF birds decreased not earlier than 20 yrs after isolation. In a region with longer and more intensive human impact (North Häme) the number of OGF species was clearly lower than in an area with more recent history of extensive forestry (eastern Kainuu). Further conservation biology projects include e.g. a survey of moths in large pristine forests of Russian Karelia and in a network of natural-like forests in Kainuu, in order to assess whether Russian OGF specialist moth populations are potential sources for Finnish populations. In a project with Mervi Koskela (see p. 40) we focus on the spatial scales at which the occurrence of six endangered spruce-log-dependent species is best explained. In a study with Karoliina Koskinen we have been censusing population densities of the moth Xestia speciosa and its larval hymenopteran parasitoids in OGF fragments of differing levels of spatiotemporal isolation. A study launched in 2003 focuses on fungal and insect communities colonizing logs of spruce, pine, birch and aspen. My aim is to map the structure of these communities, including fungi, beetles, dipterans and parasitoid wasps, and the changes of these communities along with the decaying succession of the logs during the next 10 years.
The endangered saproxylic beetle Pytho kolwensis is associated with large spruce logs. Photo by Gergely Várkonyi. Personnel - 59 Anu Väisänen, Research secretary
My background is in freshwater ecology but for the last 4 years I have been working as a research secretary mostly dealing with the administrational issues. The main tasks are to help with co- ordination of the projects, manage the budget and the payment transactions and to help to prepare proposals and reports. Daily co-ordination includes updating the web-pages and correspondence with the researches. I started my maternity leave in October 2002 and spent a year in Australia. I will return to the MRG in February 2004.
Pinnacles desert from Western Australia. Photo by Anu Väisänen. 60 - Personnel
Rongjiang Wang, Post doc
From 2002, a new collaboration project on Melitaea cinxia was started in both Finland and China. I am in charge of the work in China. Several habitats available for M. cinxia have been found in the Tianshan Mountains, Xinjiang. Similar to M. cinxia in Finland, the host plant of M. cinxia in Xinjiang is Veronica spicata. Different to the fragmented habitat patches in Åland Island, there is a large and continuous habitat for M. cinxia in Xinjiang. Therefore, it is an interesting issue to compare the features of M. cinxia between Åland and Xinjiang. We are focusing on two specific questions. The first one is whether emigration rate may be increased due to fragmentation of habitat. Mark-release-recapture experiments will be conducted on both metapopulation of M. cinxia in fragmented landscape in Åland and population in homogeneous habitat in Xinjiang. Another one is to check the genetic load and inbreeding depression of M. cinxia in both Åland and Xinjiang. Theory predicts that greater genetic load and inbreeding depression should be found in large populations living in an extensive homogeneous landscape than in the fragmented landscape. Unfortunately, because of the unexpected SARS epidemic in China in the spring of 2003, the field work in Xinjiang had to be cancelled. Without field work in the first half of this year, I was focusing on analysis of the MRR data set obtained from two species of checkerspot butterfly, Euphydryas aurinia and Melitaea phoebe, in Yanjiaping northwest of Beijing. The Virtual Migration model was used to estimate the parameters of migration. The results showed that M. phoebe had higher emigration propensity and greater daily average distance moved than those of E. aurinia, which is consistent with the genetic results that genetic differentiation was found among local populations of E. aurinia, but not in M. phoebe. In August, I came to Finland and joined the MRG. I started to develop molecular markers for identification of parentage of M. cinxia. Although the parents of almost all larval groups were clearly recorded in the cage study in Åland in the summer of 2003, the fathers of some larval groups and both parents of others were unknown yet. It is necessary to use molecular markers to identify the unknown parents. Although microsatellite DNA markers are effective for identification, I hope to develop more molecular markers in one or fewer PCRs. Now I am trying two methods, DALP (direct amplification of length polymorphisms) and AFLP (amplified fragments length polymorphisms). Facilities - 61 Laboratory facilities
Helsinki, Viikki
The MRG is located at the 5th floor of Biocenter 3, Viikinkaari 1. We have eight offices, which are currently shared by 22 people. One small office is available for minor meetings. We share a molecular laboratory with other members of the Department. The molecular laboratory is fully equipped for PCR-based projects, such as mtDNA sequencing and the screening of DNA microsatellites. Equipment includes PCR machines and centrifuges, allozyme, agarose and acrylamide gel electrophoresis equipment and facilities, an ABI 377 DNA sequencer and a MegaBace 1000 Capillary electrophoresis machine, with attached robot (both with a full-time technician who runs the machine). We also have facilities for both DNA cloning and the use of radioisotopes.
Biocenter 3. Photo by Tapio Gustafsson. 62 - Facilities
The table below shows the amount of space occupied by the MRG.
Space no. m² Office 8 166.0 Meeting room 1 8.5 DNA-laboratory*1 299.0 Insect rooms 3 52.0
* = Our share is 16%, as the laboratory is shared with the rest of the Department of Ecology and Systematics.
Rearing facilities for butterflies
In 1996 we established an insect breeding facility at the Tvärminne Zoological Station, to support experimental studies on the Glanville fritillary butterfly. From the beginning of this year the project has continued in the Viikki Biocenter, were we have separate rearing facilities for larvae and for adult butterflies. Adult butterflies are mated in a flying room with several broad spectral quartz glass fluorescent lamps. The plants needed for larval food and female egg laying are grown in a separate greenhouse, which enables the rearing of larvae all year round.
Biocenter 3 is part of The Viikki Science Park and is located 10 km from the centre of Helsinki. Field sites - 63 Field sites
Åland Islands
In the Åland Islands in SW Finland, the research is focused on the Glanville fritillary (Melitaea cinxia), its parasitoid wasps and larval host plants. The main Åland Island contains thousands of habitat patches (dry meadows) where at least one of the two larval host plant species, Plantago lanceolata and Veronica spicata, occurs, and which are considered to be potentially suitable for the Glanville fritillary. Each autumn, all known habitat patches are surveyed for the presence of larval groups of the Glanville fritillary, as well as scored for several habitat patch attributes. The latter include information on the occurrence of the host plants, landscape structure and habitat quality. In the following spring, all patches occupied in the previous autumn are re-visited, and the numbers of larval groups, numbers of larvae and numbers of cocoons of the local specialist parasitoid wasp Cotesia melitaearum are counted. This large-scale monitoring work has accumulated a comprehensive and unique database on the occurrence of the Glanville fritillary and on the habitat patch characteristics. Moreover, all the habitat patches are currently being positioned with an accurate GPS-machine, which allows analyses of the spatial patterns and properties of the patch network with GIS. Based on this monitoring work, we know the locations of the vast majority of the suitable habitat patches in Åland, totalling about 4200 patches on the main islands in 1999. The habitat patches are distributed across the Åland’s land area of 1480 km2, but most of them occur in clusters, typically of tens of patches. The main reason for the aggregated distribution of habitat patches is that the meadows mainly occur within the cultural landscape on rocky outcrops, typically near small villages. The patch- aggregates form well over one hundred habitat patch networks, many of which are occupied by the Glanville fritillary, and where the respective metapopulations have relatively independent dynamics. In 2003, about 455 patches were occupied by the Glanville fritillary.
Kuhmo
In the past 100 years, modern forestry has dramatically changed the Finnish landscape. Today the amount of old-growth forest remaining increases from the most densely populated south to the north, as well as from the western coast to the Russian border. On the other hand, because of climatic reasons, the natural species diversity of forest- specializing organisms decreases to the north. From these facts one may conclude that the most favourable area for studying forest-dwelling species is Kuhmo in eastern 64 - Field sites middle Finland. A further important feature is that there is a large extent of virgin forests nearby in Russia, which maintains viable populations of taiga species and may serve as a source for isolated Finnish populations.
Old growth forest in Jauhovaara, Kuhmo. Photo by Leena Suvanto.
These reasons led to cooperation between Ilkka Hanski’s MRG (Helsinki) and the Research Centre of the Friendship Park (Kuhmo), in the framework of a major mutual project entitled ”Biodiversity of Boreal Forests”. Research in Kuhmo has concentrated on spatial and temporal responses of old-growth forest species to forest management, on the role of two deciduous tree species in old-growth forests, and on the biology of endangered taiga species. By studying species diversity in small fragments of old-growth forest we attempt to assess the extinction debt in small forest fragments. Empirical data on a wide range of taxa, sampled from 23 forest fragments will be related to the history of forest fragmentation around each study fragment. Fragmentation may also cause extinction cascades, when the base of a specialized food chain disappears, as we have shown in the case study on a polypore fungus – moth – parasitoid chain. Aspen and goat willow, both living and dead, represent essential microhabitats for a diverse group of organisms in boreal forests. We have mapped all aspen and willow trees larger than 10 cm DBH within an area of 120 km2. Studies on spatial Field sites - 65 population structure and dynamics of organisms associated with the aspen, like the lichen Lobaria pulmonaria, several mosses and beetles, have been conducted with the aid of the database on the host tree distribution. The beetle Pytho kolwensis, living in virgin spruce forests, has been one of the target species while studying the biology of endangered old-growth forest species. Special emphasis has also been put on research on spore dispersal of the endangered saproxylic fungus Phlebia centrifuga. Long-distance spore dispersal is very rare, which probably prevents patch-to-patch colonization in fragmented landscapes. We have started a population genetic study for selected old-growth forest species including the beetle P. kolwensis. We have compared the behaviour of two moth species in a mosaic of small fragments of old-growth in otherwise managed forest landscapes. In a mark-release- recapture study we tested the significance of ’ecological corridors’ and ’stepping stones’ for the moths. Both Xestia speciosa and X. rhaetica, a generalist and an old- growth forest specialist species, respectively, preferred corridors for dispersal, but were also able to cross wide sapling stands. However, they both avoided clearcuts.
Greenland
Northeast Greenland holds one of the simplest vertebrate communities on earth. Only one rodent species is present (the collared lemming), and the site thus provides an unique opportunity to investigate the predator-prey interaction hypothesised to control rodent dynamics. A five-year field program has been organised jointly between the Universities of Helsinki, Montpellier and Freiburg from 1998 to 2002 in order to parameterize and test Hanski & Korpimäki’s (1995) model for lemming cycles. Other aspects of predator-prey interactions, such as prey selection, have also been studied. The study site is about 75 km² and is located on Traill Island (72°30’ N, 24°00’ W), NE Greenland. This area, which is part of the high Arctic breeding range of collared lemming, consists of a wide open valley facing the Kong Oscar Fjord. Numerous raised beaches lie parallel to the present shoreline. Inland elevations do not exceed 250 m above sea level and in contrast to other tundra areas, lakes are scarce and small in size (< 1 ha). Summer is short with only three months without snow cover, and mean summer temperatures do not exceed 5°C. The region has a discontinuous semi-desert polar vegetation. Lemming population dynamics display typical cyclic patterns (4 years) that are also reflected in the densities of predators: arctic fox, stoat, long-tailed skua and snowy owl. 66 - Field sites
Western part of the Greenland study area at the beginning of June. Photo by Olivier Gilg.
Wattkast
In the project “Spatial ecology of arboreal insects” we examine in what ways plant- insect interactions are affected by the spatial distribution of the host plant. For this purpose, we have mapped every individual of our target plant, the English oak (Quercus robur) within a well-defined area – the island of Wattkast in Korppoo, south-western Finland. Despite its small size (about 5 km²), Wattkast provides a diverse environment for our studies. The island consists of a patchwork of forests, fields and pastures. Most of the oak trees are situated in an area of high tree density, but individual trees are scattered over the rest of the island, enabling us to identify a range of tree densities and spatial settings for our studies. Within Wattkast, the location of all oak trees more than 50 cm in height have been accurately measured with a GPS navigator. A number of tree characteristics such as growth form, diameter of the trunk and previous herbivore damage have also been estimated, and the type of the surrounding vegetation has been classified for each oak individual. The information from the oak inventory now forms a database with information about the approximately 1600 oak individuals on the island. Field sites - 67 In September we visited a selection of small-sized oaks on Wattkast to score the presence or absence and abundance of leaf-mining and galling insect species. In the future our spatially referenced oak data will serve many other purposes, including studies of host-pathogen and multitrophic interactions.
The distribution of oak trees on Wattkast.
Madagascar: Ranomafana National Park
We have initiated a project on dung beetles inhabiting native forests in Madagascar (see p. 24). The more intensive studies have been conducted so far in the Ranomafana National Park (RNP) in southeastern Madagascar. RNP was inaugurated in 1991 with the two-fold purpose of conserving the unique biodiversity of the Ranomafana ecosystem and reducing the human pressures on the protected area. Altitudes in the park range from 500 to 1500 meters. Its 43,500 hectares are home to a diverse flora and fauna. The biological richness of Ranomafana National Park is extremely high. The tree species richness of one-hectare plots at Ranomafana, 37 families and more than 100 species, are intermediate between lower elevation rain forests in Colombia, South America and lowland rain forests in Gabon, Africa. Primate species richness is among the highest in the world, with 12 species in five families, all endemic. Land snail biodiversity is among the highest in the world. As in all of Madagascar, bird species richness is exceptionally low (110 species). The combination of high primate diversity 68 - Field sites with low avian diversity has raised questions about seed-dispersal and pollination in this forest. Research on these questions has shown that pollination and seed-dispersal at Ranomafana may be very different from other rain forest sites. Soils at RNP are mostly red clay oxisols developed from parent rock with extremely low levels of nutrients - these soils are considered some of the most naturally infertile in the world. The Namorona River bisects the park and is fed by the many small streams that flow through the park and contribute to its rugged topography. The park contains lowland rain forests, cloud forests, marshes, high plateau forests, and sections that were selectively logged before the area was protected. Mount Maharira in the southern part of the park has an unusual and rare mountain top flora and fauna shared only with a site 130 km south (Andringtra). RNP is located in the eastern moist forest climate and is hot and subtropical. November through March is considered to be the rainy season. Although the dry season extends from April to December, the North East and Central East regions have no months (or weeks) that are completely without rain. Reasonably dry months are May, September, October, and November. Torrential rain and cyclones can occur in January, February, and March. Annual rainfall ranges from 1500 to 4000 mm.
View down to the rainforest from the cascades, Ranomafana National Park. Photo by Tapio Gustafsson. Synopsis - 69 Synopsis of the year 2003
Budget
Funding Source €
Academy of Finland Spatial population processes 154 500 Metapopulation Research Group 291 660 Reserve network design: ecological and economical factors 17 520 Population biology of the trembling aspen 24 400 Developing the checkerspot butterflies as a model system in population biology 50 000 STADYCS 22 500 Spatial population dynamics and adaptation in a host-herbivore system 26 900 Salaries 143 685
University of Helsinki Centre-of-excellence funding from Ministry of Education 267 000 Salary for tehnician 32 000 Spatial ecology of forest birds in managedforest landscapes in southern Finland 21 400
Ministry of Agriculture and Forestry Biodiversity in boreal forests 21 700
Private Foundations 55 000
TOTAL 1 128 265 70 - Synopsis Publications
Cabeza, M. and Moilanen, A. 2003. Site-selection algorithms and habitat loss. Conservation Biology 17, 1402-1413
Cabeza, M, Moilanen, A. and Possingham, H.P. Metapopulation dynamics and reserve network design. In: Metapopulation Ecology, Genetics, and Evolution, Hanski, I. and Gaggiotti, O. (eds.) Academic Press, in press.
Cabeza, M., Araújo, MB., Wilson, R.J, Thomas, C.D., Cowley, M and Moilanen, A. Combining probabilities of occurrence with spatial reserve design. Journal of Applied Ecology, in press.
Dreschler, M., Frank, K, Hanski, I., O’Hara, R.B. and Wissel, C., 2003. Ranking metapopulation extinction risk: from patterns in data to conservation management decisions. Ecological Applications 13, 990-998.
Eccard, J. A., Meyer, J. and Sundell, J. Mating system, space use, and circadian activity pattern of nocturnal tree rat Thallomys nigricauda. Journal of Mammalogy, 85 (2) in press.
Elphinstone, M.S., Hinten, G.N., Anderson M.J., Nock C.J. 2003. An inexpensive and high-throughput procedure to extract and purify total genomic DNA for population studies. Molecular Ecology Notes 3(2), 317-320.
Gilg, O. 2003. La fragmentation des paysages forestiers: un problème pour la conservation de la biodiversité. In : Vallauri, D. Livre Blanc sur la protection des forêts naturelles en France. p. 87-94. Paris, Tec & Doc Lavoisier.
Gilg, O., Hanski, I. and Sittler, B. 2003. Cyclic dynamics in a simple vertebrate predator-prey community. Science 302, 866-868.
Haikola, S. 2003. Effects of inbreeding in the Glanville fritillary butterfly (Melitaea cinxia). Annales Zoologici Fennici 40, 483-493.
Hanski, I. 2003. Biology of extinctions in butterfly metapopulations. In: C. Boggs, W. Watt and P. Ehrlich (Eds.), Biology of butterflies. Synopsis - 71 Hanski, I. 2003. Ekologinen arvio Suomen metsien suojelutarpeesta. Kirjassa Harkki, S., Savola, K. ja Walsh, M. (Toim.), Palaako elävä metsä, sivut 18-33. Birdlife Suomen julkaisuja No. 5, Helsinki.
Hanski, I. 2003. Täpläverkkoperhonen metapopulaatiobiologisen tutkimuksen mallisysteeminä. Acad. Scient. Fennica, Year Book 2002.
Hanski, I. and Heino, M. 2003. Metapopulation-level adaptation of insect host plant preference and extinction–colonization dynamics in heterogeneous landscapes. Theoretical Population Biology 64, 281-290.
Hanski, I. and Ovaskainen, O. 2003. Metapopulation theory for fragmented landscapes. Theoretical Population Biology 64, 119-127.
Hinten, G.N., Harriss, F., Rossetto, M., and Baverstock, P.R. 2003. Genetic variation and island biogeography: Microsatellite and mitochondrial DNA variation in island populations of the Australian bush rat, Rattus fuscipes greyii. Conservation Genetics 4(6), 759-778.
Kankare, M. and Shaw, M.R. 2003. Molecular Phylogeny of Cotesia Cameron, 1891 (Insecta: Hymenoptera: Braconidae: Microgastrinae) Parasitoids Associated with Melitaeini Butterflies (Insecta: Lepidoptera: Nymphalidae: Melitaeini). Molecular Phylogenetics and Evolution, in press.
Laine, A-L. 2003. The first report of Phomopsis subordinaria in a natural population of Plantago lanceolata L. in Southwest Finland. Plant Pathology 52, 422.
Laine A-L. 2003. The unseen battle for life and death of anther-smut genotypes: Dynamics of multiple infection and within host competition by the anther-smut pathogen. Electronic commentary at update.bmn.com
Laine, A-L. 2003. The paradox of competitive coexistence - putting mutualism in a new light. Electronic commentary at update.bmn.com
Latva-Karjanmaa, T., Suvanto, L., Leinonen, K. and Rita, H. 2003. Emergence and survival of Populus tremula seedlings under varying moisture conditions. Can. J. For. Res. 33(11), 2081-2088. 72 - Synopsis Moya-Laraño, J. and Cabeza, M. 2003. Bimodality in the body size distribution of Mediterranean tarantula juveniles: Humphreys’ russian roulette revisited. Revista Ibérica de Aracnología 7, 211-219.
Nieminen, M., Suomi, J., van Nouhuys, S., Sauri, P. and Riekkola, M-L. 2003. Effect of iridoid glycoside content on oviposition host plant and parasitism in a specialist herbivore. Journal of Chemical Ecology 29, 823-844. van Nouhuys, S., Singer, M.C. and Nieminen, M. 2003. Spatial and temporal patterns of caterpillar performance and the suitability of two host plant species. Ecological Entomology 28, 193-202.
Ovaskainen, O. 2003. Habitat destruction, habitat restoration and eigenvector- eigenvalue relations. Mathematical Biosciences 181, 165-176.
Ovaskainen, O. and Cornell, S. 2003. Biased movement at a boundary and conditional occupancy times for diffusion processes. Journal of Applied Probability 40, 557-580.
Ovaskainen, O. and Grenfell, B. 2003. Mathematical tools for planning effective intervention scenarios for sexually transmitted diseases. Sexually Transmitted Diseases 30, 388-394.
Ovaskainen, O. and Hanski, I. 2003. How much does an individual habitat fragment contribute to metapopulation dynamics and persistence? Theoretical Population Biology 64, 481-495.
Ovaskainen, O. and Hanski, I. 2003. The species-area relationship derived from species-specific incidence functions. Ecology Letters 6, 903-909.
Ovaskainen, O. and Hanski, I. 2003. Extinction threshold in metapopulation models. Annales Zoologici Fennici 40, 81-97.
Pykälä, J., Pöyry, J., Kuussaari, M. and Heikkinen, R. 2003: Perinnebiotooppien kasvi- ja eläinlajisto. In: Tiainen, J., Kuussaari, M., Laurila, I. and Toivonen, T. (eds.): Suomen maatalousympäristön monimuotoisuus. Edita, Helsinki. In press.
Pöyry, J., Heliölä, J., Ryttäri, T. ja Alanen, A. 2003: Perinnebiotooppien lajiston uhanalaistuminen. In: Tiainen, J., Kuussaari, M., Laurila, I. ja Toivonen, T. (eds.): Suomen maatalousympäristön monimuotoisuus. Edita, Helsinki. In press. Synopsis - 73 Roslin, T. 2003. Not so quiet on the high frontier. Trends in Ecology and Evolution 18, 376-379.
Roslin, T. 2003. Lots of nurture with your nature: summer daughters make better grandmothers. Electronic commentary at update.bmn.com
Roslin, T. 2003. American coots keep count of their own eggs, even when the neighbour slinks in a few extras. Electronic commentary at update.bmn.com
Roslin, T. 2003. Mating with strangers provides no apparent fitness benefits for the coal tit. Electronic commentary at update.bmn.com
Sittler, B. and Gilg, O. 2003. Responses of Snowy Owls to lemming fluctuations: Insights from a long-term study in high arctic Greenland. Book of abstract. International Symposium: Ecology and Conservation of European Owls. 23-26 October 2003. Dornbirn (Austria).
Sittler, B. and Gilg, O. 2003. Lemming cycles in high arctic Greenland: patterns and issues emerging from an ongoing long-term study. In. 21. Internationale Polartagung der Deutschen Gesellschaft für Polarforschung. 17.-22. März 2003 Kiel. Kiel: Alfred Wegener Institut; p. 51.
Sundell, J. 2003. Lumikko myyräkantojen muokkaajana (with English summary: Effect of least weasel predation on vole population dynamics). Suomen Riista 49, 55-69.
Sundell, J. 2003. Population dynamics of microtine rodents: an experimental test of the predation hypothesis. Oikos 101, 416-427.
Sundell, J. 2003. Reproduction of the least weasel (Mustela nivalis nivalis): basic observations and the influence of the food availability. Acta Theriologica 48(1), 59- 72.
Sundell, J. 2003. Small mammal research and teaching at the Lammi Biological Station. In: From parsonage to centre of international science – half-centennial history of Lammi Biological Station (eds. Keskitalo, J. & Syrjämäki, J.). pp. 294-299. Yliopistopaino, Helsinki. (in Finnish)
Sundell, J. 2003. Ilkka Hanski, at the top of population biological research. In: From parsonage to centre of international science – half-centennial history of Lammi 74 - Synopsis Biological Station (eds. Keskitalo, J. & Syrjämäki, J.). pp. 301-306. Yliopistopaino, Helsinki. (in Finnish)
Sundell, J., Dudek, D., Klemme, I., Koivisto, E., Pusenius, J. and Ylönen, H. Variation in predation risk and vole feeding behaviour: a field test of the risk allocation hypothesis. Oecologia, in press
Sundell, J., Eccard, J., Tiilikainen, R. and Ylönen, H. 2003. Predation rate, prey preference and predator switching: experiments on voles and weasels. Oikos 101, 615-623.
Sundell, J., Huitu, O., Henttonen, H., Kaikusalo, A., Korpimäki, E., Pietiäinen, H., Saurola, P. and Hanski, I. Large-scale spatial dynamics of vole populations in Finland revealed by the breeding success of vole-eating avian predators. Journal of Animal Ecology, in press
Suvanto, L., Stenvall, N., Vares, A. and Pulkkinen, P. Alien genes in forestry: hybrid aspen as a model species for alien gene flow. Rit Mogilsar Rannsöknastöovar Skograektar, in press.
Suvanto, L., Stenvall, N., Vares, A. and Pulkkinen, P. Hybridihaavan geenivirta. Finnish Forest Research Institute Research Papers, in press.
Várkonyi, G. 2003. Population biology of periodic Xestia moths. Academic dissertation. University of Helsinki.
Várkonyi, G., Hanski, I., Rost, M. and Itämies, J. 2003. Periodic boreal moths: Population dynamic consequences of interaction with natural enemies. Entomophagous Insects Workshop. 2003. - Abstracts from the XIII International Entomophagous Insects Workshop. 30pp. Journal of Insect Science 3:33. Abstract available online: http://insectscience.org/3.33/
Várkonyi, G., Kuussaari, M. and Lappalainen, H. 2003. Use of forest corridors by boreal Xestia moths. Oecologia 137, 466-474.
Wiegand, T., Jeltsch, F., Hanski, I. and Grimm, V. 2003. Using pattern-oriented modeling for revealing hidden information: a key for reconciling ecological theory and application. Oikos 100, 209-222. Synopsis - 75 Ylönen, H., Sundell, J., Tiilikainen, R., Eccard, J. A. and Horne, T. 2003. Weasels’ (Mustela nivalis nivalis) preference for olfactory cues of the vole (Clethrionomys glareolus). Ecology 84(6), 1447-1452.
Ylönen, H., Eccard, J. A. and Sundell, J. 2003. Boreal vole cycles and vole life histories. In: Singleton, G. R., Hinds, L. A., Krebs, C. J. and Spratt, D. M. (eds). Rats, Mice and People: Rodent Biology and Management. pp. 137-141. ACIAR Monograph No. 96.
Theses
Mar Cabeza, PhD: Spatial population dynamics in reserve-network design
During the last two decades there has been a major development of methods for systematic conservation planning. The major focus has been on developing algorithms for solving problems such as which reserves should be chosen to include all species e.g. at least once. This is the so called species representation approach. More recently, the problems addressed have included some criteria for biodiversity persistence, for instance, by setting reasonable targets for species viability or by emphasizing the selection of ‘best quality sites’. However, the effect of spatial population dynamics together with the spatial configuration of reserves on species persistence has mostly been overlooked. My thesis shows that if spatial population dynamics are ignored extinctions are expected in the reserves, and that the expected proportion of extinctions is larger if habitat is modified around the reserves (e.g. unselected sites are lost). The thesis presents new methods for selecting reserve networks, that deal with spatial dynamics and habitat loss in different ways. These methods differ in complexity, going from (1) methods that qualitatively aim at achieving sets of clustered reserves; to (2) methods that use habitat models together with species-specific measures of habitat connectivity for selecting reserves according to the predicted species occurrences; to (3) methods that explicitly maximize species persistence when it is assessed with a metapopulation model that explicitly simulates the spatial dynamics of the species. The applicability of the different methods is determined by the availability of data: the more complex the model, the more information is needed.
Sari Haikola, PhD: Inbreeding depression in the Glanville Fritillary Butterfly (Melitaea cinxia)
The aim of this study was to examine the existence and severity of inbreeding 76 - Synopsis depression in the Glanville fritillary butterfly, Melitaea cinxia, metapopulation in the Åland Islands in SW Finland. The research was carried out under laboratory conditions. Controlled matings were conducted between siblings and unrelated individuals. Inbreeding is expected to be frequent in small local populations of M. cinxia in Åland. Therefore, selection during continuous inbreeding could in theory have purged the genetic load due to deleterious recessive alleles from the metapopulation. However, this study shows that even though purging has to occurred some extent, part of the genetic load still remains in the system. In the laboratory, several different traits at various life stages show inbreeding depression. These traits include important fitness components such as egg hatching rate and larval survival. The results show that inbreeding depression is manifested over the entire life-cycle of M. cinxia. As there seems to be no discrimination against close relatives as mates, inbreeding depression is expected to be an influential factor also in natural populations. In an earlier study it was found that increased homozygosity, an indication of inbreeding, was associated with elevated extinction risk of small natural populations of M. cinxia in the Åland metapopulation. The increased extinction risk of inbred M. cinxia populations was later confirmed experimentally. This study has clarified the relationship between homozygosity and the extinction risk of small natural populations.
Gergely Várkonyi, PhD: Population biology of periodic Xestia moths
Alternate-year flight of the moths in the subgenus Pachnobia (Lepidoptera, Noctuidae, Xestia spp.) in boreal forests of the Northern Hemisphere has been a population dynamic puzzle for more than a century now. The basis for periodic occurrence in Xestia is a fixed two-year life cycle and permanently dissimilar cohort sizes in most studied localities. Coexisting Xestia moths typically share common flight years, and local synchronic population dynamics tend to form very large spatial domains of odd- or even-year flight pattern. The important population dynamic question is what maintains the abundance difference between the coexisting but temporally isolated cohorts. Results from the analysis of a 22-year time series of Xestia moths and the parasitoid wasp Ophion luteus from eastern Finnish Lapland suggest that periodicity in the moth is maintained by interaction with the parasitoid. Biological and modelling results further support this hypothesis. Other natural enemies like Sorex shrews and the parasitoid Meloboris collector are likely to contribute to the regulation of minor Xestia cohorts. The transition zone between opposite-phase spatial domains of periodic occurrence in Finnish Lapland is found to be very distinct and only 10-20 km wide. Densities of both the moth and its parasitoid were very low in this zone, a result which is consistent with the predictions of the model. Extensive open areas are likely to form a dispersal barrier for Xestia and to pin the domain boundary in central Lapland. In a recently fragmented forest landscape in middle Finland movements of Synopsis - 77 adult Xestia moths tended to be canalised through old-growth forest strips (corridors). Movements of male and female moths were consistent with the pheromone-mediated mate-location syndrome. Foraging strategies and movement patterns of the old-growth spruce forest specialist X. fennica and the habitat generalist X. speciosa suggested different reproductive startegies in the two species. The extent of genetic differentiation between populations and among coexisting cohorts of X. tecta in Finnish Lapland is found to be very low, which may be due to substantial gene flow or, more likely, due to repeated population bottlenecks during the post-glacial expansion of the species from refugia in the Far East.
Johanna Ehrnsten, MSc: The within-patch movement behaviour of a mobile parasitoid, Hyposoter horticola
Many butterfly species have greatly declined worldwide, which means that their specialist parasitoids must be even more endangered. Hyposoter horticola is a very mobile, successful, specialist parasitoid of the endangered butterfly, Melitaea cinxia on the Åland islands. It finds almost all its host populations and constantly parasitises 20-30 % of its host larvae, despite the fact that parasitism is only possible during a short time period before the larvae are ready to hatch from the eggs, that is a couple of hours. It is quite exceptional for a parasitoid to be so consistent temporally and spatially usually populations fluctuate in time and space. I investigated the within-patch movement behaviour of H. horticola to see if it is possible to study the behaviour of such a mobile parasitoid in the field. I also wanted to find out how much the parasitoid moves within its habitat patches. Furthermore I wanted to find out if H. horticola has got a spatial memory and therefore is so successful in parasitising its host. The results of my experiment showed that it is possible to study the behaviour of H. horticola in the field. It is a very mobile parasitoid and it also moves considerable distances within habitat patches, probably more than its host does. The results also suggest that H. horticola learns the locations of its host´s egg clusters when they are not yet ready to be parasitised and uses that information later to visit them multiple times and parasitise them when they are ready. Another explanation is that H. horticola uses a long-range odour mark, although long-range odour marks in parasitoids are often water soluble and ephemeral and distances between egg clusters are long.
Jenni Hottola, MSc: Relationships between the structure of polypore communities and coarse woody debris in the forests of eastern Finland and adjacent areas in Russian Karelia 78 - Synopsis Wood-rotting fungi are especially susceptible to alterations in forests due to their dependence on coarse woody debris and their often specific requirements regarding microclimatic conditions and the quality of the growing substrate. Temporal and spatial continuity in the amount of suitable substrate and in environmental conditions may be critical for the survival of species and populations in the long term. Many species of wood-rotting fungi have declined in Fennoscandia during the recent decades, mainly because of forestry. In my thesis I studied the community structure of polypores (Polyporaceae s. lato) and some other species of Aphyllophorales in relation to the characteristics of coarse woody debris, and, qualitatively, also to the structure of the forest landscape and the naturalness of the stand. Particularly, I focused on rare, indicator, and red- listed species. Altogether, 37 study plots were surveyed quantitatively on both sides of the border between Finland and Russia. The results suggest that the volume of fallen dead wood is the most important factor affecting the structure of the polypore community. Also, the diversity as well as the number, diameter, and decay stage of dead trees seemed to have an effect on the incidence of species. Large stands surrounded by relatively natural forest landscapes were richer in rare and red-listed species and in the total species richness than small stands in more fragmented landscape. Rare and threatened species were most abundant in natural stands, although light thinning in the forest history didn’t seem to have a substantial impact on the incidence of polypore species. However, the latter may reflect the time delay preceding local extinctions.
Hanna Paulomäki, MSc: The direct and indirect effects of weather on the local populations of the Glanville fritillary
The aim of my master’s thesis was to study the direct and indirect effects of weather on the local populations of the Glanville fritillary in four geographically separate regions. In each region the growth of the larval groups was followed by photographing the groups on a millimetre paper. Besides this the quantity and quality of the food plants Plantago lanceolata and Veronica spicata were studied. Adult butterflies were observed using the mark-recapture method. Temperature was recorded once every five minutes on each study patch. Daily rainfall (mm/km²) was counted from the weather radar data (Finnish Meteorological Institute). There was no significant difference in average temperatures and in rainfall. Instead there was a difference in average minimum temperatures and that may have affected the growth of the larval groups. The average size of the larvae differed significantly between the four regions. In total 10 % of all egg clusters was most likely destroyed by heavy rainfall. Moreover rain destroyed the nests of some early instar larvae. Weather factors affecting the Synopsis - 79 whole island in a similar manner may cause great changes in local populations during some years because of the spatial and temporal variation in growth and development rate of the larvae and adult butterflies.
Astrid van Teeffelen, MSc: Habitat loss, connectivity and probabilities: comparing reserve selection strategies
During my graduate studies (Environmental Sciences) at Wageningen University, the Netherlands, I performed my thesis in the MRG from September 2002 until March 2003, under the supervision of Atte Moilanen. Reserve design aims at the long-term persistence of the species in the reserve network. However, most site-selection algorithms focus on species representation, but do not take species’ persistence into account. New methods have been developed in the MRG, which take species’ persistence more explicitly into account. In my thesis I have compared the results of basic reserve selection methods with the new reserve selection methods, in their ability to represent the species at a given target level. I tested the methods with a dataset on seven bird species, occurring in a study area in the Netherlands. For these birds I predicted their occurrence in the study area using various habitat models, which were based on habitat suitability, and some were extended with (species-specific) connectivity measures. By including connectivity measures, it was possible to account for habitat loss outside the reserve network, in the calculation of probabilities of occurrence. All solutions from the reserve selection methods were evaluated by assuming complete habitat loss outside the reserve network. Following, probabilities of occurrence were calculated, which were then only based on the habitat and connectivity of the reserve network itself. The results showed that basic selection methods, based on presence-absence data, select reserves that appear to be too scattered and too small to represent the species at least for a given target. The best results were obtained with methods that accounted for habitat loss outside the reserve network and that calculated probabilities of occurrence based on species-specific connectivity measures. As a consequence, these reserve networks were larger and more clustered, minimising the effect of habitat loss outside the reserve network.
Kata Valosaari, Msc: Spatial and temporal variation in mortality in the juvenile stages of the Glanville fritillary butterfly (Melitaea cinxia) in Åland islands.
The metapopulation biology of the Glanville fritillary butterfly is well known, but no local population life table study has been done before. Extinction or colonisation of local populations form the basis of metapopulation dynamics. To predict large scale metapopulation dynamics, it is important to understand also the local population 80 - Synopsis processes which may cause extinction. The aim of the my thesis was to find the key stages and the key factors for four butterfly metapopulations with life table analysis. The study metapopulations were located in different parts of the islands, in Eckerö, Finström, Jomala and Föglö. Winter mortality was the key stage of the juvenile stages in all studied metapopulations in 2000-01. About 80 % of the larvae died during winter diapause. The main cause of the high mortality is unclear but there are a few factors that increased mortality; small body size of the larvae, small larval group size, poor or late winter nest building capacity and low food plant abundance before diapause. The results of the life table analyse showed that although the mortality of the larvae varied between metapopulations in some larval instars, the total mortality did not vary between metapopulations. The total mortality of the larvae was about 95 %. When comparing the two spring larval generations separately in each metapopulation I found variation in mortality in Eckerö; mortality was higher in the spring of 2001 than in the spring of 2000. The cause of the higher morality was the poorer nutritional condition in the year 2000. Though winter mortality was the key stage of the Glanville fritillary butterfly juvenile stages in years 2000-01 generation, it may fluctuate strongly from year to year, probably depending on the weather conditions. The previous studies also shows that the mortality of any developmental stage of the butterfly fluctuates between years. The reason for this fluctuation is thought to be the weather conditions and their effect on the food plants, but also natural enemies. My results showed that predation might be important at the egg and small larval stages, but it needs further examinations to see if that’s really the case. To see if winter mortality really is the key stage of the Glanville fritillary butterfly, the life table study should be continued several years in several metapopulations.
External visits
Mar Cabeza visited Dr. M.B. Araújo, Girona (Spain), March 2003.
Mar Cabeza visited Dr. P.Williams in Natural History Museum, London (UK), March 2003.
Mar Cabeza visited the Department of Universities, Research and Information Society, Generalitat de Catalunya (Barcelona, Spain), December 2003 Synopsis - 81 Ilkka Hanski led an expedition of nine MRG members to the Ranomafana National Park in Madagascar from November 18 until December 6 (participants: Ilkka Hanski, Tapio Gustafsson, Marjo Saastamoinen, Anna-Liisa Laine, Tomas Roslin, Otso Ovaskainen, Gavin Hinten, Olivier Gilg, Gergely Várkonyi, Helena Koivulehto ja Heidi Viljanen)
Gavin Hinten visited Dr. Alfried Vogler’s lab at the Natural History Museum of London to collaborate on dung beetle phylogenetics and attended the conference on Phylogenetics and Conservation at the Royal London Zoo, February 2003.
Jonna Katajisto and Atte Moilanen visited Prof. Pierre Taberlet in Laboratoire de Biologie des Populations d’Altitude, in the University of Joseph Fourier in Grenoble, France, 21 - 26 May 2003.
Jonna Katajisto visited Prof. Jon Swenson in Department of Ecology and Natural Resource Management, in the Agricultural University of Norway, Ås, 19 - 23 October 2003.
Saskya van Nouhuys visited the Department of Multitrophic Interactions, Centre for Terrestrial Ecology, Netherlands Institute of Ecology, 1 week, March 2003.
Seminars, lectures, and talks
Mar Cabeza gave a talk “Habitat loss and connectivity in reserve network design” in Student Conference on Conservation Science, Cambridge (UK).
Mar Cabeza gave a talk “How to select reserves for biodiversity persistence?” at the LUOVA day, University of Helsinki (Finland).
Ilkka Hanski gave an invited talk in the 50th anniversity meeting of the Ecology Society of Japan. 21 March 2003.
Ilkka Hanski gave an invited lecture in the University of Sapporo in Japan. 24 March 2003.
Ilkka Hanski gave an invited talk in the annual meeting of the German ecological society in Halle. 9 September 2003 82 - Synopsis Ilkka Hanski gave an invited lecture in the meeting organized by the BirdLife International in Latvia. 10 October 2003.
Ilkka Hanski gave an invited lecture at the department of Biology, University of Haren, the Netherlands. 7 November 2003.
Ilkka Hanski talked about forest biodiversity in several national public meetings in 2003.
Paavo Hellstedt gave a talk ‘The survival rate and behaviour of captive-born released least weasel’ Paavo Hellstedt & Eva Kallio in 3rd European Congress of Mammalogy, 27 July -1 August 2003. Brno, Czech Republic.
Gavin Hinten gave a talk “Patterns and Processes of microsatellite evolution in island populations of the bush rat”. University of Helsinki lecture series, January 2003.
Gavin Hinten gave a talk “Molecular Phylogeny of Madagascar’s dung beetles”. Centre ValBio, Ranomofana National Park, December 2003.
Jenni Hottola attended the “Spatial statistics workshop” organized by Environment Graduate School, University of Oulu. Perämeri Research Station, Hailuoto, 16 - 18 October 2003.
Jonna Katajisto gave a talk “Habitat selection of Scandinavian brown bears” in Spring Symposium at the Department of Ecology and Systematics, University of Helsinki, Finland. 11 April 2003.
Jonna Katajisto gave a talk “Brown bear distribution in relation to habitats and humans” in Luova Days, University of Helsinki, Finland, 15 October 2003.
Anna-Liisa Laine gave a talk “Spatial variation in resistance in a host-pathogen system” in Spring Symposium, University of Helsinki, Finland 7 - 9 April 2003.
Anna-Liisa Laine presented a poster “ Resistance variation in a natural host-pathogen metapopulation” in ESEB IX, Leeds, UK 18 - 23 August 2003.
Saskya van Nouhuys gave a talk “Butterfly oviposition choice: Why that plant?.” 27th annual Ecology and Evolutionary Biology Symposium, Cornell University, Ithaca, New York, USA. January 2003. Synopsis - 83 Saskya van Nouhuys gave an invited talk “Mechanisms behind the population dynamics of the parasitoids of the Glanville fritillary butterfly.” Symposium on Metapopulation Biology – Achievements and Challenges, University of Helsinki, Finland. February 2003.
Saskya van Nouhuys gave an invited talk “A host and two parasitoids that each experience the same habitat differently.” Symposium on Habitat Fragmentation and Trophic Interactions in Wageningen, The Netherlands. March 2003
Saskya van Nouhuys gave an invited talk “A host and two parasitoids that each experience the same habitat differently.” Biology Department at Leiden University, The Netherlands. March 2003.
Saskya van Nouhuys gave a talk “Spatial learning (or something) allows a parasitoid to use a host that is available for an extremely short time.” XIII International Entomophagous Insect workshop in Tucson, Arizona, USA, July 2003
Saskya van Nouhuys gave a talk “Effects of iridoid glycosides on the development of herbivores and their endoparasitoids.” Annual meeting of the Entomological Society of America, Cincinnati, Ohio. USA. J. Harvey (presenter), A. Biere and S. van Nouhuys. October 2003.
Saskya van Nouhuys gave an invited talk “Spatial population dynamics in insect communities: butterflies and their host plants and parasitoids.” Department of Ecology and Evolutionary Biology seminar series at Cornell University, Ithaca, New York, USA. 10th November 2003.
Otso Ovaskainen gave a talk “From individual behavior to ecological and evolutionary metapopulation dynamics” in a seminar series of Department of Mathematics, University of Turku, Finland, 27 October 2003.
Otso Ovaskainen gave a talk “Elinympäristöt pirstoutuvat – miten populaatiot selviytyvät?” at the monthly meeting of Suomen biologinen seura Vanamo, 27 October 2003.
Otso Ovaskainen gave a talk “Metapopulation dynamics in fragmented landscapes” in the Symposium “Metapopulation biology – achievements and challenges” organized by the Department of Ecology and Systematics, University of Helsinki, Finland, 14 February 2003. 84 - Synopsis Otso Ovaskainen gave a talk “Dispersal in ecological (and evolutionary) landscapes” in a seminar series of Department of Ecology and Systematics, University of Helsinki, Finland, 5 February 2003.
Reijo Penttilä gave a talk “Effects of habitat loss and fragmentation on old-growth polypore species” in a seminar “Metapopulation Biology – Achievements and Challenges” held in University of Helsinki, Viikki, 14 - 15 February 2003.
Reijo Penttilä gave a talk “Kulotusalan kääväkkäät” in a seminar “Metsänkehitys palaa alkuun – 10 vuotta Kitsin palosta” held in University of Joensuu, 10 - 11 June 2003.
Reijo Penttilä gave a talk “ Kääpien lajimäärät talous- ja luonnonmetsissä” and attended to a panel conversation in a meeting “Kääpien inventointi luonnon- ja talousmetsissä – tiedettä ja viestintää” arranged by Metsäbiologian kerho and held in Tieteiden talo, Helsinki, 7 November 2003.
Tomas Roslin gave the talk “Äkämähyönteiset metsärengaskehrääjän runtelemissa kanadalaismetsissä” [Galling insects in the feet of ravaging forest tent caterpillars.] The Entomological Society of Finland; Helsinki, Finland, 11 April 2003.
Alia Sarhan gave a talk “Parental relatedness and offspring fitness” in Spring Symposium, University of Helsinki, Finland 7 - 9 April 2003.
Janne Sundell gave a talk “Large-scale spatial dynamics of vole populations in Finland revealed by the breeding success of vole-eating avian predators” in 4th European Congress of Mammalogy, Brno, Czech Republic, July 2003.
Leena Suvanto gave a talk “Alien genes in forestry: hybrid aspen as a model species for alien gene flow” at “Forest Genetic Resources: Their Use and Conservation” meeting in Húsavík, Iceland. 27 - 30 August 2003.
Leena Suvanto gave an invited talk “Hybridihaavan geenivirta” in Finnish Forest Research Institute’s Punkaharju research station seminar in Vantaa, Finland. 13 November 2003.
Gergely Várkonyi gave a talk “Periodisten harmoyökkösten ekologia” in the Finnish Entomological Society, 21 March 2003. Synopsis - 85 Gergely Várkonyi gave a talk “Periodic boreal moths: population dynamic consequences of interaction with natural enemies” in the XIII International Entomophagous Insect Workshop in Tucson AZ, 27 - 31 July 2003.
TV, radio, and newspapers
Review of the papers (provisional) more than 200 reports from more than 25 countries)
Cyclic Dynamics in a Simple Vertebrate Predator-Prey Community Olivier Gilg, Ilkka Hanski, and Benoît Sittler Science 302 (Oct 31 2003): 866-868
Radio – TV: BBC World (International): Interview for the News (live). Oct. 31 CNN International (International): Dec. 14 TV5 Europe (Europe): Interview for the News. Nov. 17 CNN (USA): « next@cnn ». Nov. 22. Discovery Channel (USA). Oct. 31 Channel 4 (UK): Interview for the News (live). Oct. 31 CBC (Canada): Interview for “As It Happens”. Oct. 31 ABC (Australia). Oct. 31 France 2 (France): Interview for the News. Nov. 17 Finnish Radio (Finland): Interview for the News. Oct. 30 West Deutche Rundfunk (Germany) Südwestrundfunk 2 (Germany): Interview. Oct. 31. Südwestrundfunk (Germany): Interview. Nov. 5. Polskie Radio (Poland): Oct. 31., Etc.
Newspapers: The Inter. Herald Tribune (Internat.): Nov. 4. The Holland Sentinel (USA) The New York Times (USA): Oct. 31. Orlando Sentinel (USA): Nov. 9 The Washington Post (USA) Die Welt (Germany): Nov. 2 (Top Stories, front page) The Times (UK): Oct. 31 Stern (Germany) (Top Stories, front page) Frankfurter Allgem. Zeitung (Germany): Nov. 1. The Guardian (UK): Oct. 31. Tagesspiegel (Germany): Nov. 2/3. The Independent (UK): Oct. 31. Die Welt am Sontag (Germany): Nov. 11. The Globe and Mail (Canada): Oct. 31. Badische Zeitung (Germany): Oct. 31. The Halifax Tribune (Canada): Nov. 2. Süddeutsche Zeitung (Germany): Oct. 31. The Australian (Australia): Nov. 8. Dresdner Neueste Nachr. (Germany): Oct. 31. New Scientist: Nov. Westdeutsche Zeitung (Germany): Oct. 31. Le Monde (France): Nov. 7. Marburger Anzeiger (Germany): Oct.31. Libération (France): Nov. 6. Südkurier (Germany): Oct. 31. Le Figaro (France) : Nov. 1. Offenburger Tagblatt (Germany): Oct.31. Science et Vie (France) : Feb. 2004 Bergedorfer Zeitung (Germany): Oct.31. Science et avenir (France): Nov. Donau Kurier (Germany): Oct. 31. La Croix (France): Nov. Stuttgarter Zeitung (Germany): Oct. 31. Le Bien Public (France) : Nov. 19. Hamburger Abendblatt (Germany): Oct. 31. Le Journal de Saône et Loire (France): Nov. 11. Zürichsee-Zeitungen (Switzerland): Nov. 8. La Gazette de Montpellier (France): Nov. 21. Basler Zeitung (Switzerland): Nov. 3. Revue Nationale de la Chasse (France) : Feb. Zurcher Zeitung (Switzerland): Nov. Dagens Nyheter (Sweden): Oct. 31. Politiken (Denmark): Nov. 9. San Diego Union Tribune (USA): Oct. 31. Hindustan Times (India) Pittsburgh Post-Gazette (USA): Oct. 31. Taipei Times (Taiwan): Nov. 2. San Fransisco Times (USA) Helsingin Sanomat (Finland). Nov. 18. Salt Lake Tribune (USA): Nov. 6. Yliopistolehti (Finland): Nov. Star Tribune (USA): Oct. 31. Atuagagdliutit/Grønlandsposten (Greenland): Dec. 4. The Buffalo News (USA): Nov. 9. + Dutch, Austrian Newspapers, Etc.
Press Agencies: UPI - United Press International. Finnish Press Agency (Finland). Oct. 30. AFP - French Press Agency (France). Oct. 30. Schweitzeriche Press Argentur (Swiss). Oct. 31 DPA - Deutsche Press Argentur (Germany). Etc. 86 - Synopsis Oliver Gilg was interviewed for several TV programs (BBC World, CNN International, TV5 Europe, CNN, Discovery Channel, Channel 4, CBC, ABC, France 2, etc.) on “Cyclic dynamics in a simple vertebrate predator-prey community” published in Science 302 (Oct 31 2003).
Oliver Gilg was interviewed for several Newspapers (The Inter. Herald Tribune, The New York Times, The Washington Post, The Times, The Guardian, The Independent, The Globe and Mail, The Australian, New Scientist, Le Monde, Die Welt, Stern, etc.) on “Cyclic dynamics in a simple vertebrate predator-prey community” published in Science 302 (Oct 31 2003).
Niclas Fritzén was interviewed on the Melitaea cinxia metapopulation project on the Åland Islands for a TV program (YLE2, FST Uteliv, ed. Per-Ole Svarvar), 10 October 2003.
Ilkka Hanski was interviewed for a TV program on forest conservation in September 2003, for a radio science program on a paper on lemming dynamics published in Science in November 2003, and for a major newspaper on forest conservation in November 2003.
Jonna Katajisto was interviewed on brown bear research in YLE1 Aamutelevisio, “Karhun jäljillä”, 31 October 2003.
Jonna Katajisto and Janne Sundell were interviewed on brown bear research, “Karhun käyttäytyminen kiinnostaa” Ympäristö-lehti (ed. Leena Huttunen), August 2003.
Mikko Kuussaari, Juha Pykälä and Juha Pöyry were interviewed on the effects of grassland management on different taxa, “Niityt tarvitsevat yksilöllistä hoitoa”, Helsingin Sanomat, Science pages (ed. Arja Kivipelto), 12 July 2003.
Juha Pöyry was interviewed on the effects of grassland management on butterflies and moths, “Yli sadan perhosen elämä on riippuvainen maataloudesta”, Luomu, Nr. 6 (ed. Brita Suokas), October 2003.
Tomas Roslin was interviewed on dung beetle ecology for the programme Vihreä Laakso (Salon seudun paikalliskanava), August 2003.
Tomas Roslin was interviewed on dung beetle ecology for the magazine Luonnonvara. Synopsis - 87 Marjo Saastamoinen, Niclas Fritzén, Anne Holma and Kata Valosaari were interviewd on the Melitaea cinxia dispersal experiment carried out in Sund, Åland, “De lever fjärilsliv i Sund”, Nya Åland, 11 June 2003.
Teaching and courses
523052 Conservation Biology in Fragmented Landscapes, 2-4 stw.
This course will give information on the spatial structure and dynamics of fragmented populations. The topics of the lectures deal with habitat loss, fragmentation and species extinction; metapopulation concepts and modelling; the causes and patterns of extinctions; dispersal in fragmented landscapes; population genetics of fragmented populations and the selection of natural reserves. Empirical case studies of the Melitaea cinxia metapopulation in Åland islands will be presented. The lectures are given by the researchers of the Metapopulation Research Group.
The course consists of:
- lectures from Tue 23rd Sept until 30th October (1 sw, obligatory) - computer exercises on metapopulation dynamics (1sw, optional) - seminar based on given litterature in Lammi biological station on Fri 31st Oct - Sun 2nd Nov (1sw, obligatory) - book exam based on Hanski, I. (1999) Metapopulation Ecology. Oxford University Press, Oxford.(1sw, optional).
The lectures:
1. Introduction to metapopulations: Habitat loss, fragmentation and species extinction. Atte Moilanen
2. Extinctions: causes and patterns. Atte Moilanen
3. Dispersal, connectivity and colonisation. Atte Moilanen
4. Putting it all together: Spatially realistic metapopulation modelling. Atte Moilanen
5. Habitat loss, the extinction debt and the delay to extinction. Otso Ovaskainen 88 - Synopsis 6. Population genetics of fragmented landscapes I. Gavin Hinten
7. Population genetics of fragmented landscapes II. Gavin Hinten
8. Empirical case study: Cinxia metapopulation in Åland I. Ilkka Hanski 9. Empirical case study: Cinxia metapopulation in Åland II. Ilkka Hanski
10. Design of reserve networks I: Introduction. Mar Cabeza
11. Design of reserve networks II: Site selection algorithms. Mar Cabeza
12. Conclusions and prospects for the future. Ilkka Hanski
Other teaching
Mar Cabeza taught at the international course on “Plant dispersal”, Oulanka field station, Kuusamo, Finland.
Paavo Hellstedt gave a lecture in the course of “Animal identification”. Dept. of Ecology & Systematics, University of Helsinki.
Paavo Hellstedt served as a leading teacher at the field course “Ecology”. Dept. of Ecology & Systematics, University of Helsinki.
Paavo Hellstedt served as a teacher on the field course and seminar “Research planning and methods” in Kilpisjärvi. Dept. of Ecology & Systematics, University of Helsinki.
Gavin Hinten gave a lecture in the post-graduate course: “Molecular methods in ecology”. University of Helsinki, September 2003.
Gavin Hinten gave four lectures in the undergraduate course: “Laboratory methods in ecological genetics”. University of Helsinki, November 2003.
Jenni Hottola was a teacher in the “Polypore course” organized by the Finnish Nature League in Nuuksio, 23 - 25 May 2003.
Jonna Katajisto gave lectures about bear biology and conservation “Suurpetoleiri” organized by Kainuun luonnonsuojelupiiri in Kuhmo, 5 - 7 September 2003. Synopsis - 89 Jonna Katajisto gave a lecture in course “Ecological applications of geographic information systems”, 25 Nov 2003.
Anna-Liisa Laine gave a lecture “Ecosystem processes and hierarchy – implications for conservation of species“ in the course “From individuals to ecosystems”.
Atte Moilanen and Jonna Katajisto gave a course in “Programming ecological simulations”, 1 - 8 September 2003.
Tomas Roslin gave a lecture in the course “Ecological Genetics” (coordinated by Oscar Gaggiotti, Juha Merilä and Craig Primmer).
Gergely Várkonyi taught on the field course on “Taxonomy and Ecology of Terrestrial Animals”, University of Oulu, Oulanka Biological Station, Finland, 5 - 14 August 2003.
Honors and awards
Ilkka Hanski was invited to become a Honorary Professor in the Peking Normal University, China, in 2003
Jenni Hottola was elected the MSc of the Year in Department of Biology, University of Oulu, Finland.
Anna-Liisa Laine was awarded Olli’s prize for the best student presentation at the Spring Symposium for PhD students of Department of Ecology and Systematics, University of Helsinki, Finland.
Council Memberships
Ilkka Hanski has served in the Scientific Advisory Board of the University of Helsinki. Ilkka Hanski has been a board member of the Finnish Cultural Foundation.
Ilkka Hanski has been a board member of the E.J. Sariola Foundation, Finland.
Marko Nieminen has served as the vice chairman of conservation committee of the Finnish Lepidopterological Society. 90 - Synopsis Juha Pöyry has served as a secretary of the Consultative Committee for Butterfly and Moth Conservation in Finland.
Juha Pöyry has served as a member of the Conservation Committee of the Lepidopterological Society of Finland.
Juha Pöyry has served as a member of the Conservation Committee of threatened Hymenoptera in Finland. Juha Pöyry has served as the secretary of the Research programme for biodiversity in agricultural environments, coordinated by the Ministry of Agriculture and Forestry. Tomas Roslin has served as a member of the board of the Societas Entomologica Helsingforsiensis.
Gergely Várkonyi has served as a member of the Committee for Hymenopteran Conservation (Finnish Ministry of the Environment).
Meetings organized by the MRG
Metapopulation Biology - Achievements and Challenges, 14 - 15 February 2003.
Visitors to the MRG
Prof. Mark Burgman, University of Melbourne, Australia. Examiner to Mar Cabeza´s academic dissertation “Spatial population dynamics in reserve-network design” and collaborative research, 23 - 30 April 2003.
Prof. Jon Swenson, the Department of Biology and Nature Conservation, Agricultural University of Norway, discuss collaboration with Scandinavian Brown Bear research Project, 7 - 11 April 2003.
Dr. Juan Corley, National Scientific Research Council of Argentina, Laboratory of Insect Ecology, Bariloche, Argentina. Collaborative research on parasitoid population ecology, 21 June - 4 July 2003.
Dr. C.J. Nagelkerke, Institute for Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam, The Netherlands. Collaborative research, 7 - 21 September 2003. Synopsis - 91 Mr. Chandrakant Maheswari, the Indian Institute of Technology, Delhi, India, worked as a research assistant in the Metapopulation Research Group, 1 May - 3 August 2003.
Dr. Brendan Wintle, University of Melbourne, Australia. Collaboration concerning habitat modelling and reserve selection, 3 - 6 November 2003.
Vice President Mr Li Zhuqi, Director General Mr Han Jianguo, Director General Mr Jing Daping, Director General Mr Peng Lianming, Director of Division Mr Tang Xianming, Director of Division Mr Lu Rongkai and Professor Mr He Minghong, National Natural Science Foundation of China (NSFC). Excursion to one of the centres-of-excellence in research, 25 August 2003.
A typical Melitaea cinxia habitat patch in the Åland Islands. Photo by Marko Nieminen. 92 - Prospects Prospects for the year 2004
Many of us look forward to some celebration in the early 2004 to mark the publication of the two edited volumes that have taken so much of our time in the past two years, On the Wings of Checkerspots (edited by Paul Ehrlich and Ilkka Hanski, published by the Oxford University Press) and the Ecology, Genetics, and Evolution of Metapopulations (edited by Ilkka Hanski and Oscar Gaggiotti, published by the Academic Press). In terms of research projects, the checkerspot project in China should eventually get started, having been delayed by the SARS epidemics in 2003; field work on dung beetles in Madagascar will continue, and perhaps another MRG expedition to Madagascar will take place in the autumn; and the cage experiments on the Glanville fritillary will continue in Åland and expand to the Tvärminne Zoological Station. Two new projects will address the 4th goal that we have set to our research, to disseminate our research results beyond the scientific community. A teaching package including a web site will be prepared for primary and secondary schools. This material is essentially based on our research on the Glanville fritillary and it will first be offered to the schools in Åland, as a small sign of appreciation of all the help and support that we have received in Åland over the past decade. Second, Hanski will coordinate a discussion forum for scientists working with questions related to forests and forestry, with the aim of encouraging analytic debate on scientific issues that too often remain buried under non-scientific arguments. We expect to see substantial turnover of students in 2004, and we will have several long-term visitors. We are approaching the end of the current Centre of Excellence funding period, with more assessment of our performance and achievements. The future of the Centre of Excellence programme is still unclear, but it is likely that preparations for the new round will start in 2004.
Ilkka Hanski