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Department of Geosciences and Geography
Physical Geography

Miska Luoto

Selected publications 2005-2016


1. Refereed international journal articles


  1. Riihimäki, H., Heiskanen, J. & Luoto, M. (2016) The effect of topography on arctic-alpine aboveground biomass and NDVI patterns. International Journal of Applied Earth Observations and Geoinformation (in press).
  2. Taka, M., Aalto, J. & Luoto, M. (2016) The direct and indirect effects of watershed land use and soil type on stream water metal concentrations. Water Resources Research (in press).
  3. Niskanen, A., Heikkinen, R.K., Väre, H. & Luoto, M. (2016) Improving forecasts of arctic-alpine refugia persistence with landscape-scale variables. Geografiska Annaler: Series A, Physical Geography (in press).
  4. Niskanen, A., Luoto, M. , Väre, H. & Heikkinen, R.K. (2016) Models of Arctic-alpine refugia highlight importance of climate and local topography. Polar Biology (in press - online).
  5. Väliranta, M., Salojärvi, N., Vuorsalo, A., Juutinen, S., Korhola, A, Luoto, M. & Tuittila, E-S. (2016) Holocene fen-bog transitions, current status in Finland and future perspectives. Holocene (in press - online).
  6. Mod, H.K., Scherrer, D., Luoto, M. & Guisan, A. (2016) What we use is not what we know: environmental predictors in plant distribution models. Journal of Vegetation Science (in press - online).
  7. Mateo, R.G., Broennimann, O., Normand, S., Petitpierre, B., Araújo, M.B., Svenning, J-C., Baselga, A., Fernández-González, F., Gómez-Rubio, V., Muñoz, J., Suarez, G.M., Luoto, M. , Guisan, A., Vanderpoorten, A. (2016) The mossy north: an inverse latitudinal diversity gradient in European bryophytes. Scientific Reports 6, 25546.
  8. Mod, H., Heikkinen, R.K., le Roux, P.C., Wisz, M.S. & Luoto, M. (2016) Impact of biotic interactions on biodiversity varies across a landscape. Journal of Biogeography (in press).
  9. Pajunen, V., Luoto, M. & Soininen, J. (2016) Stream diatom assemblages as predictors of climate. Freshwater Biology 61, 876-886.
  10. Stewart, L., Alsos, I.G., Bay, C., Breen, A.L., Brochmann, C., Boulanger-Lapointe, N., Daniëls, F.J., Broennimann, O., Bültmann, H., Bøcher, P.K., Damgaard, C., Daniëls1, F.J.A., Ehrich, D., Eidesen, P.B., Guisan, A., Jónsdóttir, I.S., Lenoir, J., le Roux, P.C., Lévesque, E., Luoto, M., Nabe-Nielsen, J., Schönswetter, P., Tribsch, A., Tveraabak, L.U., Virtanen, R., Walker, D.A., Westergaard, K.B., Yoccoz, N.G., Svenning, J-C, Wisz M., Schmidt, N.M., & Pellissier, L. (2016). The regional species richness and genetic diversity of Arctic vegetation reflect both past glaciations and current climate. Global Ecology and Biogeography 25, 430–442.
  11. Salonen, J.S., Verster, A.J., Engels, S., Soininen, J., Trachsel, M. & Luoto, M. (2016) Calibrating aquatic microfossil proxies with regression-tree ensembles: cross-validation with modern chironomid and diatom data. Holocene (in press).
  12. Pellissier, L., Eidesen, P. B., Ehrich, D., Descombes, P., Schönswetter, P., Tribsch, A., Westergaard, B.K., Alvarez, N., Guisan, A., Zimmermann,NE, Normand, S., Vittoz, P., Luoto, M., Damgaard, C., Brochmann, C., Wisz, M.S. & Alsos, I. G. (2016). Past climate-driven range shifts and population genetic diversity in arctic plants. Journal of Biogeography 43, 461–470.
  13. Mod, H., Heikkinen, R.K., le Roux, P.C., Väre, H. & Luoto, M. (2016) Contrasting effects of biotic interactions on richness and distribution of vascular plants, bryophytes and lichens in an arctic-alpine landscape. Polar Biology 39, 649-657.
  14. Wasof, S., Lenoir, J., Aarrestad, P.A., Alsos, I.G., Armbruster, W.S., Austrheim, G., Bakkestuen, V., Birks, H.J.B., Bråthen, K.A., Broennimann, O., Brunet, J., Bruun, H.H., Dahlberg, C.J., Dieckmann, M., Dullinger, S., Dynesius, M., Ejrnæs, R., Gégout, J.-C., Graae, B.J., Grytnes, J.A., Guisan, A., Hylander, K., Jónsdóttir, I.S., Kapfer, J., Klanderud, K., Luoto, M. , Milbau, A., Moora, M., Nygaard, B., Odland, A., Pauli, H., Ravolainen, V., Reinhardt, S., Sandvik, S.M., Schei, F.H., Speed, J., Svenning, J.-C., Thuiller, W., Tveraabak, L.U., Vandvik, V., Velle, L.G., Virtanen, R., Vittoz, P., Willner, W., Wohlgemuth, T., Zimmermann, N.E., Zobel, M., Decocq, G. (2016) Disjoint populations of European vascular plant species keep the same climatic niches. Global Ecology and Biogeography 24, 1401-1412.
  15. Nylén, T. & Luoto, M. (2016) Influence of patch size and connectivity on beach and dune species in land uplift coasts. Plant Ecology & Diversity 9, 35-44.
  16. Pajunen, V., Luoto, M. & Soininen, J. (2016) Climate is an important driver for stream diatom distributions. Global Ecology and Biogeography 25, 198-206.


  1. Heino, J., Erkinaro, J., Huusko, A., & Luoto, M. (2015). Climate change effects on freshwater fishes, conservation and management. Conservation of Freshwater Fishes, 76. Cambridge University Press.
  2. Tallavaaraa, M., Luoto, M. , Korhonen, N., Järvinen, H., & Seppä, H. (2015) Human population dynamics in Europe over the Last Glacial Maximum. Proceedings of the National Academy of Sciences 112, 8232-8237.
  3. Taka, M., Aalto, J. & Luoto, M. (2015) Spatial modelling of stream water quality along the urban-rural gradient. Geografiska Annaler: Series A, Physical Geography 97, 819-834.
  4. Kuusijärvi, A., Hardwick, B., Luoto, M. & Roslin, T. (2015) A stable, genetically determined colour dimorphism in the dung beetle Aphodius depressus: patterns and mechanisms. Ecological Entomology 40, 575-584.
  5. Kuussaari, M., Heikkinen, R.K., Heliölä, J., Luoto, M. , Mayer, M., Rytteri, S., & von Bagh, P. (2015) Successful translocation of the threatened Clouded Apollo butterfly (Parnassius mnemosyne) and metapopulation establishment in southern Finland. Biological Conservation 190, 51-59.
  6. Mod, H., le Roux, P.C., Guisan, A. & Luoto, M. (2015) Biotic interactions boost spatial models of species richness. Ecography 38, 913–921.
  7. Varanka, S., Hjort, J., & Luoto, M. (2015) Geomorphological factors predict water quality in boreal rivers. Earth Surface Processes and Landforms 40, 1989-1999.
  8. Nylén, T., Hellemaa, P. & Luoto, M. (2015) Determinants of sediment properties and organic matter in beach and dune environments based on boosted regression trees. Earth Surface Processes and Landforms 40, 1137–1145.
  9. Hylander, K., Ehrlén, J., Luoto, M. & Meineri, E.. (2015) Microrefugia: Not for everyone. AMBIO 44, 60-68
  10. Soininen, J., Bartels, P., Heino, J., Luoto, M. & Hillebrand, H. (2015) Towards more integrated ecosystem research in aquatic and terrestrial environments. BioScience 65, 174-182.
  11. Meier, S., Luoto, M. , & Soininen, J. (2015) The effects of local, buffer zone and geographical variables on lake plankton metacommunities. Hydrobiologia 743, 175-188.
  12. Nylen, T. & Luoto, M. (2015) Primary succession, disturbance and productivity drive complex species richness patterns on land uplift beaches. Journal of Vegetation Science 26, 267-277.


  1. Soininen, J. & Luoto, M. (2014) Predictability in species distributions: a global analysis across organisms and ecosystems. Global Ecology and Biogeography 23, 1264-1274.
  2. Aalto, J., Venäläinen, A., Heikkinen, R.K. & Luoto, M. (2014) Potential for extreme loss in high-latitude Earth surface processes due to climate change. Geophysical Research Letters 41, 3914–3924.
  3. Suvanto, S., le Roux, P.C. & Luoto, M. (2014) Arctic-alpine vegetation biomass is driven by fine scale abiotic heterogeneity. Geografiska Annaler: Series A, Physical Geography 96, 549-560.
  4. le Roux, P.C., Pellissier, L., Wisz, M.S. & Luoto, M. (2014) Incorporating dominant species as proxies for biotic interactions strengthens plant community models. Journal of Ecology 102, 767–775.
  5. Aalto, J. & Luoto, M. (2014) Integrating climate and local factors for geomorphological distribution models. Earth Surface Processes and Landforms 39, 1729-1740.
  6. Salonen, J.S., Luoto, M. , Alenius, T., Heikkilä, M., Seppä, H., Telford, R. & Birks J. (2014) Reconstructing palaeoclimatic variables from fossil pollen using boosted regression trees: comparison and synthesis with other quantitative reconstruction methods. Quaternary Science Reviews 88, 69-81.
  7. Mod, H., le Roux, P.C. & Luoto, M. (2014) Outcomes of biotic interactions are dependent on multiple environmental variables. Journal of Vegetation Science 25, 1024–1032.
  8. Aalto, J., le Roux, P.C., & Luoto, M. (2014) The meso-scale drivers of temperature extremes in high-latitude Fennoscandia. Climate Dynamics 42, 237–252.
  9. le Roux, P.C. & Luoto, M. (2014) Earth surface processes drive the richness, composition and occurrence of plant species in an arctic-alpine environment. Journal of Vegetation Science 25, 45-54.


  1. Aalto, J., le Roux, P.C. & Luoto, M. (2013) Vegetation mediates soil temperature and moisture in arctic-alpine environments. Arctic, Antarctic, and Alpine Research 45, 429-439.
  2. Nylen, T., le Roux, P.C. & Luoto, M. (2013) Biotic interactions drive species occurrence and richness in dynamic beach environments. Plant Ecology 214, 1455-1466.
  3. le Roux, P.C., Aalto, J. & Luoto, M. (2013) Soil moisture’s underestimated role in climate change impact modelling in low energy systems. Global Change Biology 19, 2965–2975.
  4. Blank, L., Luoto, M. & Merilä, J. (2013) Potential effects of climate change on the distribution of the common frog Rana temporaria at its northern range margin. Israel Journal of Ecology & Evolution 59, 130–140.
  5. Parviainen, M., Zimmermann, N.E., Heikkinen, R.K. & Luoto, M. (2013) Using unclassified continuous remote sensing data to improve distribution models of red-listed plant species. Biodiversity and Conservation 22, 1731–1754.
  6. Eskildsen, A., le Roux, P.C., Heikkinen, R.K., Høye, T.T., Kissling, W.D., Pöyry, J., Wisz, M.S. & Luoto, M. (2013) Testing species distribution models across space and time: high latitude butterflies and recent warming. Global Ecology and Biogeography 22, 1293-1303.
  7. Hjort, J. & Luoto, M. (2013). Statistical methods for geomorphic distribution modeling. In: Shroder, J. (Editor in Chief), Baas, A.C.W. (Ed.). Treatise on Geomorphology. Academic Press, San Diego, CA, vol. 2, : Quantitative Modeling of Geomorphology, pp. 59-73.
  8. Walker, D.A., Alsos, I.G., Bay, C., Boulanger-Lapointe, N., Breen, Al.L., Bültmann, H., Christensen, T., Damgaard, C., Daniëls, F.J.A., Hennekens, S., Raynolds, M.K., le Roux, P.C., Luoto, M. , Pellissier, L., Peet, R.K., Schmidt, N.M., Stewart, L., Virtanen, R., Yoccoz, N.G. & Wisz, M.S. (2013) Rescuing valuable Arctic vegetation data for biodiversity models, ecosystem models and a Panarctic vegetation classification. Arctic 66, 134-137.
  9. Lenoir, J., Graae, B.J., Aarrestad, P.A., Alsos, I.G., Armbruster, W.S., Austrheim, G., Bergendorff, C., Birks, H.J.B., Bråthen, K.A., Brunet, J., Bruun, H.H., Dahlberg, C.J., Decocq, G., Diekmann, M., Dynesius, M., Ejrnæs, R., Grytnes, J-A., Hylander, K., Klanderud, K., Luoto, M., Milbau, A., Moora, M., Nygaard, B., Odland, A., Ravolainen, V.T., Reinhardt, S., Sandvik, S.M., Schei, F.H., Speed, J.D.M., Tveraabak, L.U., Vandvik, V., Velle, L.G., Virtanen, R., Zobel, M., Svenning J-C. Local temperatures inferred from plant communities suggest strong spatial buffering of climate warming across Northern Europe. Global Change Biology 19, 1470–1481.
  10. le Roux, P.C., Virtanen, R., & Luoto, M. (2013) Geomorphological disturbance is necessary for predicting fine-scale species distributions. Ecography 36, 800–808.
  11. le Roux, P.C., Lenoir, J., Pellissier, L., Wisz, M.S. & Luoto, M. (2013) Horizontal, but not vertical, biotic interactions affect fine-scale plant distribution patterns in a low energy system. Ecology 94, 671-682.
  12. Soininen, J., Korhonen, J. & Luoto, M. (2013) Stochastic species distributions are driven by organism size. Ecology 94, 660-670.
  13. Virtanen, R., Grytnes, J-A., Lenoir, J., Luoto, M., Oksanen, J., Oksanen, L. & Svenning, J-C. (2013) Productivity-diversity patterns in arctic tundra vegetation. Ecography 36, 331–341.
  14. Wisz, M.S., Pottier, J., Kissling, W.D., Pellissier, L., Lenoir, J., Damgaard, C.F., Dormann, C.F., Forchhamer, M.C., Grytnes, J.A., Guisan, A., Heikkinen, R.K., Høye, T.T., Kühn, I., Luoto, M., Maiorano, L., Nilsson, M.C., Normand, S., Öckinger, E., Schmidt, N.M., Termansen, M., Timmermann, A., Wardle, D.A., Aastrup, P., & Svenning, J.C. (2013) The role of biotic interactions in shaping distributions and realised assemblages of species: implications for species distribution modelling. Biological Reviews 88, 15-30.


  1. le Roux, P.C., Virtanen, R., Heikkinen, R.K. & Luoto, M. (2012) Biotic interactions affect the elevational ranges of high-latitude plant species. Ecography 35, 1048–1056.
  2. Hyvönen, T., Luoto, M. & Uotila, P. (2012) Assessment of weed establishment risk in a changing European climate. Agricultural and Food Science 21, 348-360.
  3. Lenoir, J., Virtanen, R., Oksanen, J., Oksanen, L., Luoto, M. , Grytnes, J.-A., & Svenning, J.-C. (2012) Dispersal ability links to cross-scale species diversity patterns across the Eurasian Arctic tundra. Global Ecology and Biogeography 21, 851-860.
  4. Hjort, M., Heikkinen R.K. & Luoto, M. (2013) Inclusion of explicit measures of geodiversity improve biodiversity models in a boreal landscape. Biodiversity and Conservation 21, 3487–3506.
  5. Varanka, S. & Luoto, M. (2012) Environmental determinants of water quality in boreal rivers based on partitioning methods. River Research and Applications 28, 1034–1046.
  6. Salonen, J.S., Seppä, H., Luoto, M. , Bjune, A.E., & Birks J. (2012) A North European pollen-climate calibration set: analysing the climatic responses of a biological proxy using novel regression tree methods. Quaternary Science Reviews 45, 95-110.
  7. Hjort, M. & Luoto, M. (2012) Can geodiversity be predicted from space? Geomorphology 153-154, 74–80.
  8. Soininen, J. & Luoto, M. (2012) Is catchment productivity a useful predictor of taxa richness in lake plankton communities? Ecological Applications 22, 624-633.
  9. Heikkinen, R.K., Marmion, M. & Luoto, M. (2012) Does the interpolation accuracy of species distribution models come at the expense of transferability? Ecography 35, 276–288.
  10. Astorga, A., Oksanen, J., Luoto, M. , Soininen, J., Virtanen, R. & Muotka, T. (2012) Distance decay of similarity in freshwater communities: do macro- and microorganisms follow the same rules? Global Ecology and Biogeography 21, 365–375.


  1. Astorga, A., Heino, J., Luoto, M. & Muotka, T. (2011) Freshwater biodiversity at regional extent: determinants of macroinvertebrate taxonomic richness in headwater streams. Ecography 34, 705-713.
  2. Fronzek, S.,Carter, T.R. & Luoto, M. (2011) Evaluating sources of uncertainty in modelling the impact of probabilistic climate change on sub-arctic palsa mires.Natural Hazards and Earth System Sciences 11, 2981–2995.
  3. Alahuhta, J., Vuori, K.M. & Luoto, M. (2011) Land use, geomorphology and climate as environmental determinants of littoral helophyte occurrence and percentage cover in boreal catchments. Boreal Environment Research 16, 185-202.
  4. Sormunen, H., Virtanen, R. & Luoto, M. (2011) Inclusion of local environmental conditions alters high-latitude vegetation change predictions based on bioclimatic models. Polar Biology 34, 883–897.
  5. Hjort, M. & Luoto, M. (2011) Novel theoretical insights into geomorphic process–environment relationships using simulated response curves. Earth Surface Processes and Landforms 36, 363-371.
  6. Alahuhta, J., Heino, J. & Luoto, M. (2011) Climate change and the future distributions of aquatic macrophytes across boreal catchments. Journal of Biogeography 38, 383-393 .


  1. Virtanen, R., Luoto, M., Rämä, T., Mikkola, K., Hjort, J.; Grytnes, J-A., & Birks, J. (2010) Recent vegetation changes in the high-latitude tree-line ecotone are controlled by geomorphologic disturbance, productivity and diversity. Global Ecology and Biogeography 19, 810-821.
  2. Rocchini, D., Balkenhol, N., Carter, G.A., Foody, G.M., Gillespie, T.W., He, K.S., Kark, S., Levin, N. Lucas, K., Luoto, M., Nagendra, H., Oldeland, J., Ricotta, C., Southworth, J., & Neteler, M. (2010) Remotely sensed spectral heterogeneity as a proxy of species diversity: recent advances and open challenges. Ecological Informatics 5, 318-329.
  3. Kangas, K. Luoto, M., Ihantola, A., Tomppo, E., & Siikamäki, P. (2010) Recreation induced changes in boreal bird communities in protected areas. Ecological Applications 20, 1775-1786.
  4. Maes, D., Titeux, N., Hortal, J., Anselin, A., Decleer, K., De Knijf, G., Fichefet, V. & Luoto, M. (2010) Predicted insect diversity declines under climate change in an already impoverished region. Journal of Insect Conservation 14, 485-489.
  5. Luoto, M., Marmion, M., & Hjort, J. (2010) Assessing the spatial uncertainty in predictive geomorphological mapping: a multi-modelling approach. Computers and Geosciences 36, 355–361.
  6. Parviainen, M., Luoto, M. & Heikkinen, R.K. (2010) NDVI -based productivity and heterogeneity as indicators of species richness in boreal landscapes. Boreal Environment Research 15, 301-318.
  7. Kaartinen, S., Luoto, M. & Kojola, I. (2010) Selection of den sites by wolves (Canis lupus) in boreal forests in Finland. Journal of Zoology 281, 99-104.
  8. Virkkala, R., Marmion, M., Heikkinen, R.K., Thuiller, W. & Luoto, M. (2010) Predicting bird species distribution patterns under the changing climate: influence of modeling technique and topography. Acta Oecologica 36, 269-381.
  9. Fronzek, S., Carter, T.R., Räisänen, J., Ruokolainen, L. & Luoto, M. (2010) Applying probabilistic projections of climate change with impact models: a case study for sub-arctic palsa mires in Fennoscandia. Climatic Change 99, 515-534.
  10. Heikkinen, R.K., Luoto, M., Leikola, N., Pöyry, J., Settele, J., Kudrna, O., Marmion, M., Fronzek, S. & Thuiller, W. (2010) Assessing the vulnerability of European butterflies to climate change using multiple criteria. Biodiversity and Conservation 19, 695–723.
  11. Hjort, M. & Luoto, M. (2010) Geodiversity of high-latitude landscapes in northern Finland. Geomorphology 115, 109-116.


  1. Marmion, M., Luoto, M., Heikkinen, R. K. & Thuiller, W. (2009) The performance of state-of-the-art modelling techniques depends on geographical distribution of species. Ecological Modelling 220, 3512-3520.
  2. Parviainen, M., Luoto, M. & Heikkinen, R.K. (2009) The role of local and landscape level productivity in modelling of boreal plant species richness. Ecological Modelling 220, 2690-2701.
  3. Kaartinen, S., Luoto, M. & Kojola, I. Carnivore-livestock conflicts: determinants of wolf (Canis lupus) depredation on sheep farms in Finland. Biodiversity and Conservation 18, 3503–3517.
  4. Hjort, M. & Luoto, M. (2009) Interaction of geomorphic and ecologic features across altitudinal zones in a subarctic landscape. Geomorphology 112, 324-333.
  5. Parviainen, M., Marmion, M., Luoto, M., Thuiller, W. & Heikkinen, R.K. (2009) Using summed individual species models and state-of-the-art modelling techniques to identify threatened plant species hotspots. Biological Conservation 2501-2509.
  6. Roslin, T., Avomaa, T., Leonard, M., Luoto, M. & Ovaskainen, O. (2009) Some like it hot: microclimatic variation affects the abundance and movements of a critically endangered dung beetle. Insect Conservation and Diversity 2, 232-241.
  7. Titeux, N., Maes, D., Marmion, M., Luoto, M., & Heikkinen, R.K. (2009) Inclusion of soil data improves the performance of bioclimatic envelope models for insect species distributions in temperate Europe. Journal of Biogeography 36, 1459-1473.
  8. Partanen S., Luoto M. & Hellsten S. (2009) Habitat level determinants of emergent macrophyte occurrence, abundance and change in two boreal lakes in Finland. Aquatic Botany 90, 261-268.
  9. Raatikainen, K., Luoto, M. & Heikkinen, R. K. (2009) Relative importance of habitat area, connectivity, management and local factors for vascular plants: spring ephemerals in boreal semi-natural grasslands. Biodiversity and Conservation 18, 1067-1085.
  10. Marmion, M., Parviainen, M. & Luoto, M., Heikkinen, R.K. & Thuiller, W. (2009) Evaluation of consensus methods in predictive species distribution modeling. Diversity and Distributions 15, 59-69.
  11. Pöyry J., Luoto M., Heikkinen R.K., Kuussaari M. & Saarinen K. (2009) Species traits explain recent range shifts of Finnish butterflies. Global Change Biology 15, 732-743.
  12. Marmion, M., Hjort, J., Thuiller, W. & Luoto, M. (2009) An improved approach for predictive geomorphological mapping: the statistical consensus method. Computers and Geosciences 35, 615-625.


  1. Marmion, M., Hjort, J., Thuiller, W. & Luoto, M. (2008) A comparison of predictive methods in modelling the distribution of periglacial landforms in Finnish Lapland. Earth Surface Processes and Landforms 33, 2241-2254.
  2. Kivinen, S., Luoto, M., Heikkinen, R.K. & Saarinen, K. (2008) Threat spots and environmental determinants of red-listed plant, butterfly and bird species in boreal agricultural environments. Biodiversity and Conservation 17, 3289-3305.
  3. Parviainen, M., Luoto, M., Ryttäri, T. & Heikkinen, R. K. (2008) Modelling the occurrence of threatened plant species in taiga landscapes: methodological and ecological perspectives. Journal of Biogeography 35, 1888-1905.
  4. Virkkala, R., Heikkinen, R.K., Leikola, N. & Luoto, M. (2008) Projected large-scale range reduction of northern-boreal land bird species due to climate change. Biological Conservation 141, 1343-1353.
  5. Pöyry, J., Luoto, M., Heikkinen, R.K. & Saarinen, K. (2008) Species traits contribute to the uncertainty of bioclimatic models. Global Ecology and Biogeography 17, 403-414.
  6. Ovaskainen, O., Luoto, M., Ikonen, I., Rekola, H., Meyke, E. & Kuussaari, M. (2008) An empirical test of a diffusion model: predicting clouded apollo movements in a novel environment. The American Naturalist 170, 610-619.
  7. Luoto, M. & Hjort, J. (2008) Downscaling of coarse-grained geomorphological data. Earth Surface Processes and Landforms 33, 75-89.
  8. Hjort, J. & Luoto, M. (2008) Can abundance of geomorphological features be predicted using presence–absence data? Earth Surface Processes and Landforms 33, 741-750.
  9. Luoto, M. & Heikkinen, R.K. (2008) Disregarding topographical heterogeneity biases species turnover assessments based on bioclimatic models. Global Change Biology 14, 483-494.
  10. Mitikka, V., Heikkinen, R.K., Luoto, M., Araújo, M.B., Saarinen, K., Pöyry, J. & Fronzek, S. (2008) Predicting range expansion of the map butterfly in Northern Europe using bioclimatic models. Biodiversity and Conservation 17, 623-641.


  1. Luoto, M., Virkkala, R., Heikkinen R.K. (2007) The role of land cover in bioclimatic models depends on spatial resolution. Global Ecology and Biogeography 16, 34-42.
  2. Piha, H., Luoto, M., Piha, M. & Merilä, M. (2007) Anuran abundance and persistence in agricultural landscapes during a climatic extreme. Global Change Biology 13, 300-311.
  3. Heikkinen, R.K., Luoto, M., Toivonen, T & Kuussaari, M. (2007) Effects of model complexity, spatial resolution and modelling technique on distribution modelling of a threatened butterfly. Landscape and Urban Planning 79, 347-357.
  4. Luoto, M. (2007) New insights into factors controlling drainage density in subarctic landscapes. Arctic, Antarctic, and Alpine Research 39, 117-126.
  5. (2007) Pelagic fish abundance in relation to regional environmental variations in the Gulf of Finland, Northern Baltic Sea. ICES Journal of Marine Science 64, 487-495.
  6. Hjort, J., Luoto, M. & Seppälä, M. (2007) Landscape scale determinants of periglacial features in subarctic Finland: a grid-based modelling approach. Permafrost and Periglacial Processes 18, 115-127.
  7. Parviainen, M. & Luoto, M. (2007) Climate envelopes of mire complex types of Fennoscandia. Geografiska Annaler A 89, 137-151.
  8. Kuussaari, M., Heliölä, J. Luoto, M. & Pöyry, J. (2007) Determinants of local species richness of butterflies and moths in boreal agricultural landscapes. Agriculture, Ecosystems & Environment 122, 366-376.
  9. Kivinen, S., Luoto, M., Kuussaari, M. & Saarinen, K. (2007) Differing effects of land cover and climate on species richness of declining, stable and increasing butterflies in boreal agricultural landscapes. Agriculture, Ecosystems & Environment 122, 453-460.
  10. Heikkinen, R.K., Luoto, M., Virkkala, R., Pearson, R. G. & Körber, J-H. (2007) Biotic interactions improve the prediction of boreal bird distributions at macro-scales. Global Ecology and Biogeography 16, 754-763.
  11. Araújo, M.B. & Luoto, M. (2007) The importance of biotic interactions for modelling species’ responses under climate change. Global Ecology and Biogeography 16, 743-753.
  12. Piha, H., Luoto, M. & Merilä, J. (2007) Amphibian occurrence is influenced by current and historic landscape characteristics. Ecological Applications 17, 2298-2309.


  1. Hjort, J. & Luoto, M. (2006) Modelling patterned ground distribution in Finnish Lapland: an integration of topographical, ground and remote sensing information. Geografiska Annaler 88 A, 19-29.
  2. Kivinen, S., Luoto, M., Kuussaari, M. & Helenius, J. (2006) Multi-species richness of boreal agricultural landscapes: effects of climate, habitat, soil and geographical location. Journal of Biogeography 33, 862-875.
  3. Partanen, S. & Luoto, M. (2006) Environmental determinants of littoral paludification in boreal lakes. Limnologica 36, 98-109.
  4. Heikkinen, R.K., Luoto, M. & Virkkala, R. (2006) Does fine-tuning of climatic predictors improve the performance of bioclimatic envelope models for migratory birds? Diversity and Distributions 12, 502-510.
  5. Fronzek, S., Luoto, M. & Carter, T.R. (2006) Potential effect of climate change on the distribution of palsa mires in subarctic Fennoscandia. Climate Research 32, 1-12.
  6. Luoto, M., Heikkinen, R.K., Pöyry, J. & Saarinen, K. (2006) Determinants of biogeographical distribution of butterflies in boreal regions. Journal of Biogeography 33, 1764-1778.
  7. Luoto, M. & Hjort, J. (2006) Scale matters – a multi-resolution study of the determinants of patterned ground activity in subarctic Finland. Geomorphology 80, 282-294.
  8. Heikkinen, R.K., Luoto, M., Araújo, M.B., Virkkala, R., Thuiller, W. & Sykes, M. (2006) Methods and uncertainties in bioclimatic envelope modelling under climate change. Progress in Physical Geography 30, 751-777.
  9. Austin, M. P., Belbin, L., Meyers, J.A., Doherty, M.D. & Luoto, M. (2006) Evaluation of statistical models for predicting plant species distributions: construction of artificial data based on ecological theory. Ecological Modelling 199, 197-216.
  10. Pöyry, J., Luoto, M., Paukkunen, J. Pykälä, J. Raatikainen, K. & Kuussaari, M. (2006) Different responses of plants and herbivore insects to a gradient of vertebrate grazing. Oikos 115, 401-412.


  1. Pykälä, J., Luoto, M., Heikkinen, R.K. & Kontula, T. (2005) Plant species richness and persistence of rare plants in abandoned semi-natural grasslands in northern Europe. Basic and Applied Ecology 6, 25-33.
  2. Luoto, M. & Hjort, J. (2005) Evaluation of current statistical approaches for predictive geomorphological mapping. Geomorphology 67, 299-315.
  3. Heikkinen, R.K., Luoto, M., Kuussaari, M. & Pöyry, J. (2005) New insights to butterfly-environment relationships with partitioning methods. Proceedings of the Royal Society of London Series B Biological Sciences 272, 2203-2210.
  4. Virkkala, R., Luoto, M., Heikkinen, R.K. & Leikola, N. (2005) Distribution patterns of boreal marshland birds: modelling the relationships to land cover and climate. Journal of Biogeography 32, 1957-1970.
  5. Luoto, M., Pöyry, J., Heikkinen, R.K. & Saarinen, K. (2005) Uncertainty of bioclimate envelope models based on geographical distribution of species. Global Ecology and Biogeography 14, 575-584.