Peer-reviewed scientific articles
2025 —————————————————————————————————————————-
Yan, X., Virtanen, R. & Eskelinen, A. 2025. Nutrient addition and herbivore exclusion alter plant traits and biomass via distinct mechanisms: intraspecific variability vs. species turnover. New Phytologist, in press.
Huovinen, L., M. Saastamoinen, J. M. Chase, A.-L. Laine, A. Räsänen, and A. Eskelinen. 2025. Herbivory Modifies the Role of Spatial Processes in a Grassland Plant Metacommunity. Ecology Letters 28, no. 11: e70257. https://doi.org/10.1111/ele.70257.
Johanson, N., Olde Venterink, H., Carvalheiro, L.G., Eskelinen, A. & Virtanen, R. 2025. Long‐term nutrient addition and grazing exclusion determine flower abundance, diversity and community composition in high‐latitude grasslands. Oikos: e11562. https://doi.org/10.1002/oik.11562
Virtanen, R., Borer, E.T., Crawley, M., Ebeling, A., Harpole, W.S., Risch, A.C., Roscher, C., Schütz, M., Seabloom, E.W. & Eskelinen, A. 2025. Neglecting non‐vascular plants leads to underestimation of grassland plant diversity loss under experimental nutrient addition. Journal of Ecology 113: 1672–1685. https://doi.org/10.1111/1365-2745.70052.
Carroll, O.H., Seabloom, E.W., Borer, E.T., Harpole, W.S., Wilfahrt, P., Arnillas, C.A., … Eskelinen, A., …, et al. 2025. Frequent failure of nutrients to increase plant biomass supports the need for precision fertilization in agriculture. Scientific Reports 15, 14564.
Barrio, I.C., Vuorinen, K.E.M., Barbero-Palacios, L., Defourneaux, M., Petit Bon, M., Greer, E.A., …, Eskelinen, A., …, et al. 2025. Emerging priorities in terrestrial herbivory research in the Arctic. Arctic Science 11: 1-16.
Chen, Q., Blowes, S.A., Harpole, W.S., Ladouceur, E., Borer, E.T., MacDougall, A., …, Eskelinen, A., …, et al. 2025. Local nutrient addition drives plant diversity losses but not biotic homogenization in global grasslands. Nature Communications: 4903.
Fay, P.A., Gherardi, L.A., Yahdjian, L., Adler, P.B., Bakker, J.D., …, Eskelinen, A., …, et al. 2025. Interactions among nutrients govern the global grassland biomass–precipitation relationship. Proceedings of the National Academy of Sciences 122: e2410748122.
Pärtel, M., Tamme, R., Carmona, C.P., Riibak, K., Moora, M., Bennett, J.A., …, Eskelinen, A., … et al. 2025. Global impoverishment of natural vegetation revealed by dark diversity. Nature 641: 917–924.
Spohn, M., Bagchi, S., Bakker, J.D., Borer, E.T., Carbutt, C., Catford, J.A., …, Eskelinen, A., … et al. 2025. Interactive and unimodal relationships between plant biomass, abiotic factors, and plant diversity in global grasslands. Communications Biology 8: 97.
Nelson, R.A., Sullivan, L.L., Hersch-Green, E.I., Seabloom, E.W., Borer, E.T., Tognetti, P.M., …, Eskelinen, A., …Johanson, N., …, et al. 2025. Forb diversity globally is harmed by nutrient enrichment but can be rescued by large mammalian herbivory. Communications Biology 8: 444.
Wei, G.-W. & van Kleunen, M. 2025. Responses of naturalized alien plants to soil heterogeneity and competition vary with the global naturalization success of the native competitors. Journal of Ecology 113: 1532–1545.
Ma, M., Eskelinen, A., Zhao, Y., Baskin, C.C., Xin, C., Zhang, P., Guo, Z., Zhang, H., Wang, X., Zhang, P., Du, G. 2025. Multiple mechanisms associated with loss of seed bank diversity under nitrogen enrichment. Journal of Ecoloy 113: 649-661.
2024 —————————————————————————————————————————-
MacDougall, A.S., Esch, E., Chen, Q., Carroll, O., Ohlert, T., Siewert, M., …, Eskelinen, A., …, et al. 2024. Widening global variability in grassland biomass since the 1980s. Nature Ecology & Evolution 8: 1877–1888.
De Carlan, C., Kaarlejärvi, E., De Tender, C., Eskelinen, A., and Verbruggen, E. 2024. Shifts in mycorrhizal types of fungi and plants in response to fertilisation, warming and herbivory in a tundra grassland. New Phytologist 243: 1190 – 1204.
Virtanen, R., Harpole, WS., Dunker, S., and Eskelinen, A. 2024. Multiple global change factors cause declines of a temperate bryophyte. Plant Ecology & Diversity: 1-12.
Fu, Q.-Y., Wei, G.-W., Wang, M.-Z., Cui, Y., Dong, B.-C. & Yu, F.-H. 2024. Contrasting responses of an invasive plant to herbivory of native and introduced insects. Ecological Processes 13: 78.
2023 —————————————————————————————————————————-
Siebert, J., Sünnemann, M., Hautier, Y., Risch, A.C., Bakker, J.D., Biederman, L., …, Eskelinen, A., …et al. 2023. Drivers of soil microbial and detritivore activity across global grasslands. Communications biology 6: 1220.
Chen, Q., Wang, S., Borer, E.T., Bakker, J.D., Seabloom, E.W., …, Eskelinen, A., … et al. 2023. Multidimensional responses of grassland stability to eutrophication. Nature Communications 14: 6375.
Garcia Criado, M., Myers-Smith, I.H., Bjorkman, A.D., Normand, S., …, Eskelinen, A., … et. al. 2023. Plant traits poorly predict winner and loser shrub species in a warming tundra biome. Nature Communications 14: 3837.
Eskelinen, A., Jessen, MT., Bahamonde, H.A., Bakker, J.D., Borer, E.T., Caldeira, M.C., et al. 2023. Herbivory and nutrients shape grassland soil seed banks. Nature Communications 14: 3949. https://doi.org/10.1038/s41467-023-39677-x
Pichon, N.A., Kaarlejärvi, E. & Eskelinen, A. 2023. Seed limitation interacts with biotic and abiotic factors to constrain novel species’ impact on community biomass and richness. Ecology Letters 26: 908-918. https://doi.org/10.1111/ele.14219
Jessen, M.-T., Auge, H., Harpole, S. & Eskelinen, A. 2023. Litter accumulation, not light limitation, drives early plant recruitment. Journal of Ecology 111: 1174-1187. https://doi.org/10.1111/1365-2745.14099
Muehleisen, A.J., Watkins, C.R.E., Altmire, G.R., Shaw, E.A., Case, M.F., …, Eskelinen, A., …, et al. 2023. Nutrient addition drives declines in grassland species richness primarily via enhanced species loss. Journal of Ecology 111: 552-563. https://doi.org/10.1111/1365-2745.14038
2022 —————————————————————————————————————————-
Beat, F., Moser, B., Tytgat, B., Zimmermann, S., Alberti, A., Biederman. L.A., …, Eskelinen, A., …, et al. 2022. Long-term N-addition alters the community structure of functionally important N-cycling soil microorganisms across global grasslands. Soil Biology and Biochemistry: 108887. https://doi.org/10.1016/j.soilbio.2022.108887
Risch, A.C., Zimmermann, S., Schütz, M., Borer, E.T., Broadbent, A.A.D., Caldeira, M.C., …, Eskelinen, A., …, et al. 2023. Drivers of the microbial metabolic quotient across global grasslands. Global Ecology & Biogeography 32: 904-918. https://doi.org/10.1111/geb.13664
Eskelinen, A., Harpole, W.S., Jessen, M.-T., Virtanen, R., & Hautier, Y. 2022. Light competition drives herbivore and nutrient effects on plant diversity. Nature 611: 301–305. DOI: 10.1038/s41586-022-05383-9
Jessen, M.-T., Auge, H., Hautier, Y., Harpole, S. & Eskelinen, A. 2022. Grazing and light modify Silene latifolia responses to nutrients and future climate. Plos One. https://doi.org/10.1371/journal.pone.0276789
Virtanen, R., Bakker, J.D., Jessen, M.-T., Sullivan, L.L., Harpole, W.S., & Eskelinen, A. 2022. Is the bryophyte soil diaspore bank buffered against nutrient enrichment and grazing exclusion? Plant and Soil 477: 487–499. https://doi.org/10.1007/s11104-022-05450-0
Vázquez, E., Schleuss, P.-M., Borer, E.T., Bugalho, M.N., Caldeira, M.C., …, Eskelinen, A., …, et al. 2022. Nitrogen but not phosphorus addition affects symbiotic N2 fixation by legumes in natural and semi-natural grasslands located on four continents. Plant and Soil 478: 689-707. https://doi.org/10.1007/s11104-022-05498-y
Cappelli, S, Pichon, N.A., Mannall, T, Allan, E. 2022. Partitioning the Effects of Plant Diversity on Ecosystem Functions at Different Trophic Levels.” Ecological Monographs 92(3): e1521. https://doi.org/10.1002/ecm.1521
Chen, Q., Wang, S., Seabloom, E., MacDougall, A., Borer, E., Bakker, J., …, Eskelinen, A., …, et al. 2022. Nutrients and herbivores impact grassland stability across spatial scales through different pathways. Global Change Biology 28: 2678-2688. https://doi.org/10.1111/gcb.16086
Langley, J.A., Grman, E., Wilcox, K.R., Avolio, M.L., Komatsu, K.J., Collins, S., …, Eskelinen, A., …, et al. 2022. Do tradeoffs govern plant species responses to different global change treatments? Ecology 103(6): e3626. https://doi.org/10.1002/ecy.3626
2021 —————————————————————————————————————————-
Pichon, N.A., Cappelli, S. L., & Allan, E. 2021. Intraspecific trait changes have large impacts on community functional composition but do not affect ecosystem function. Journal of Ecology 110: 644–658. https://doi.org/10.1111/1365-2745.13827
Walde, M., Allan, E., Cappelli, S. L., Didion‐Gency, M., Gessler, A., Lehmann, M. M., Pichon, N. A., & Grossiord, C. (2021). Both diversity and functional composition affect productivity and water use efficiency in experimental temperate grasslands. Journal of Ecology 109: 3877-3891. https://doi.org/10.1111/1365-2745.13765
Werner, C., Tuomi, M. & Eskelinen, A. 2021. Trait-based responses to cessation of nutrient enrichment in a tundra plant community. Oecologia 197: 675-684. https://doi.org/10.1007/s00442-021-05064-w
Ebeling, A., Strauss, A., Adler, P.B., Arnillas, C., Barrio, I., Biederman, L., …, Eskelinen, A., … et al. 2021. Nutrient enrichment increases invertebrate herbivory and pathogen damage in grasslands. Journal of Ecology 110:327-339. https://doi.org/10.1111/1365-2745.13801
Radujković, D., Verbruggen, E., Seabloom, E.W., Bahn, M., Biederman, L.A.,, Borer, E.T., …, Eskelinen, A., … et al. 2021. Soil properties as key predictors of global grassland production: have we overlooked micronutrients? Ecology Letters 24:2713-2725. https://doi.org/10.1111/ele.13894
Wilfahrt, P.A., Asmus, A.L., Seabloom, E.W., Henning, J.A., Adler, P., Arnillas, C.A., …, Eskelinen, A., … et al. 2021. Temporal rarity is a better predictor of local extinction risk than spatial rarity. Ecology 102(11): e03504. https://doi.org/10.1002/ecy.3504
Seabloom, E., Batzer, E., Chase, J., Harpole, S., Adler, P., Bagchi, S., …, Eskelinen, A., … et. al. 2021. Species loss due to nutrient addition increases with spatial scale in global grasslands. Ecology Letters 24: 2100–2112. https://doi.org/10.1111/ele.13838.
Eskelinen, A., Elwood, E., Harrison, S., Beyen, E. & Gremer, J. 2021. Vulnerability of grassland seed banks to resource-enhancing global changes. Ecology 102:e03512. https://doi.org/10.1002/ecy.3512
Van Sundert, K., Khan, M.A.S.A., Bharath, S., Buckley, Y.M., Caldeira, M.C., Donohue, I., …, Eskelinen, A., … et al. 2021. Fertilized graminoids intensify negative drought effects on grassland productivity. Global Change Biology 27: 2441-2457. https://doi.org/10.1111/gcb.15583
2020 —————————————————————————————————————————-
Borer, E., Harpole, W.S., Adler, P., Arnillas, C., Bugalho, M., Cadotte, M., …, Eskelinen, A., … et al. 2020. Nutrients cause grassland biomass to outpace herbivory. Nature Communications 11: 6036. https://doi.org/10.1038/s41467-020-19870-y
Hautier, Y., Zhang, P., Loreau, M., Wilcox, K.R., Seabloom, E.W., Borer, E.T., … Eskelinen, A., … et al. 2020. General destabilizing effects of eutrophication on grassland productivity at multiple spatial scales. Nature Communications 11: 5375. https://doi.org/10.1038/s41467-020-19252-4
Jessen, M.-T., Kaarlejärvi, E., Olofsson, J. & Eskelinen, A. 2020. Mammalian herbivory shapes intraspecific trait responses to warmer climate and nutrient enrichment. Global Change Biology 26: 6742-6752. https://doi.org/10.1111/gcb.15378
Risch, A., Zimmermann, S., Ochoa-Hueso, R., Schütz, M., Frey, B. …, Eskelinen, A., … et al. 2020. Global impacts of fertilization and herbivore removal on soil net nitrogen mineralization are modulated by local climate and soil properties. Global Change Biology 26: 7173-7185. https://doi.org/10.1111/gcb.15308
Eskelinen, A., Gravuer, K., Harpole, S., Harrison, S., Virtanen, R. & Hautier, Y. 2020. Resource-enhancing global changes drive a whole-ecosystem shift to faster cycling but decrease diversity. Ecology 101: e03178. https://doi.org/10.1002/ecy.3178
Ochoa Hueso, R., Borer, E.T., Seabloom, E.W., Hobbie, S.E., Risch, A.C., …, Eskelinen, A., … et al. 2020. Microbial processing of plant remains is co-limited by multiple nutrients in global grasslands. Global Change Biology 26: 4572-4582. https://doi.org/10.1111/gcb.15146
Gravuer, K., Eskelinen, A., Winbourne, J.B. & Harrison, S. 2020. Vulnerability and resistance in the spatial heterogeneity of soil microbial communities under resource additions. Proceedings of the National Academy of Sciences 117: 7263-7270. https://doi.org/10.1073/pnas.1908117117
Haydn, T., Bjorkman, A. , Isla Myers-Smith, I. , Elmendorf, S., Kattge, J., Diaz, S.,…, Eskelinen, A., … et al. 2020. Global plant trait relationships extend to the climatic extremes of the tundra biome. Nature Communications 11: 1351. https://doi.org/10.1038/s41467-020-15014-4
2019 —————————————————————————————————————————-
Risch, A., Zimmermann, S., Ochoa-Hueso, R., Schütz, M., Frey, B. …, Eskelinen, A., … et al. 2019. Soil net nitrogen mineralization across global grasslands. Nature Communications 10: 4981. https://doi.org/10.1038/s41467-019-12948-2
Komatsu, K.J, Avolio, M.L., Lemoine, N.P., Isbell, F. …, Eskelinen, A., … et al. 2019. Global change effects on plant communities are magnified by time and the number of global change factors imposed. Proceedings of the National Academy of Sciences 116: 17867 – 17873. https://doi.org/10.1073/pnas.1819027116
Thomas, H.J.D., Myers‐Smith, I.H., Bjorkman, A.D., Elmendorf, S.C. …, Eskelinen, A., … et al. 2019. Traditional plant functional groups explain variation in economic but not size‐related traits across the tundra biome. Global Ecology and Biogeography 28: 78 – 95. https://doi.org/10.1111/geb.12783
2018 —————————————————————————————————————————-
Bjorkman, A.D., Myers-Smith, I., Elmendorf S., Normand, S. Thomas, H.J.D. …, Eskelinen, A., …et al. 2018. Tundra Trait Team: A database of plant traits spanning the tundra biome. Global Ecology and Biogeography 27: 1402 – 1411. https://doi.org/10.1111/geb.12821
Bjorkman, A.D., Myers-Smith, I., Elmendorf S., Normand, S. Rüger, N. …, Eskelinen, A., …et al. 2018. Changes in plant functional traits across a warming tundra biome. Nature 562: 57 – 61. https://doi.org/10.1038/s41586-018-0563-7
Hautier, Y., Isbell, F., Borer, E.T., Seabloom, E.W., Harpole, W.S. …, Eskelinen, A., … et al. 2018. Local loss and spatial homogenization of plant diversity reduce ecosystem multifunctionality. Nature Ecology and Evolution 2: 50 – 56. https://doi.org/10.1038/s41559-017-0395-0
2017 —————————————————————————————————————————-
Kaarlejärvi, E., Eskelinen, A., & Olofsson, J. 2017. Herbivores rescue diversity in warming tundra by modulating trait-dependent species losses and gains. Nature Communications 8, art. 419. https://doi.org/10.1038/s41467-017-00554-z
Eskelinen, A., Kaarlejärvi, E. & Olofsson, J. 2017. Herbivory and nutrient limitation protect warming tundra from lowland species’ invasion and diversity loss. Global Change Biology 23: 245 – 255. https://doi.org/10.1111/gcb.13397
Wilcox, K.R., Tredennick, A.T., Koerner, S.E., Grman, E., Hallett, L.M. …, Eskelinen, A., … et al. 2017. Asynchrony among local communities stabilises ecosystem function of metacommunities. Ecology Letters 20: 1534 – 1545. https://doi.org/10.1111/ele.12861
Gravuer, K., Eskelinen, A. 2017. Nutrient and rainfall additions shift phylogenetically estimated traits of soil microbial communities. Frontiers in Microbiology 8: art. 1271. https://doi.org/10.3389/fmicb.2017.01271
2016 —————————————————————————————————————————-
Eskelinen, A., Saccone, P., Spasojevic, M. & Virtanen, R. 2016. Herbivory mediates the long-term shift in the relative importance of microsite and propagule limitation. Journal of Ecology 104: 1326 – 1333. https://doi.org/10.1111/1365-2745.12592
Copeland S., Harrison, S., Latimer, A., Damschen, E., Eskelinen, A., Fernandez-Going, B., Spasojevic, M., Anacker, B. & Thorne, J. 2016. Ecological effects of extreme drought on Californian herbaceous plant communities. Ecological Monographs 86: 295 – 311. https://doi.org/10.1002/ecm.1218
Virtanen, R., Eskelinen, A., Harrison, S. 2016. Comparing the responses of bryophytes and short-statured vascular plants to climate shifts and eutrophication. Functional Ecology 31: 946 – 954. https://doi.org/10.1111/1365-2435.12788
2015 —————————————————————————————————————————-
Eskelinen, A. & Harrison, S. 2015. Resource co-limitation governs plant community responses to altered precipitation. Proceedings of the National Academy of Sciences 112: 13009 – 13014. https://doi.org/10.1073/pnas.1508170112
Eskelinen, A. & Harrison, S. 2015. Biotic context and soil properties modulate native plant responses to enhanced rainfall. Annals of Botany 116: 963 – 973.
Harrison, S., Damschen, E., Fernandez-Going, B., Eskelinen, A. & Copeland, S. 2015. Plant communities on infertile soils are less sensitive to climate change. Annals of Botany 116: 1017 – 1022. https://doi.org/10.1093/aob/mcv109
Stark, S., Männistö, M. & Eskelinen, A. 2015.When do grazers accelerate or decelerate soil carbon and nitrogen cycling in tundra? – A test of theory on grazing effects in fertile and infertile habitats. Oikos 124: 593 – 602. https://doi.org/10.1111/oik.01355
Eskelinen, A. & Harrison, S. 2015. Erosion of beta diversity under interacting global change impacts in a semiarid grassland. Journal of Ecology 103: 397 – 407. https://doi.org/10.1111/1365-2745.12360
Virtanen, R., Eskelinen, A. and Harrison, S. 2015. Bryophyte diversity in Californian grasslands in relation to substrate quality, exotic vascular plants and disturbance. Biodiversity and Conservation 24: 103-116. https://doi.org/10.1007/s10531-014-0793-4
2014 —————————————————————————————————————————-
Saccore, P., Pyykkönen, T., Eskelinen, A. & Virtanen, R. 2014. Environmental perturbation, grazing pressure and soil wetness jointly drive mountain tundra toward divergent alternative states. Journal of Ecology 102: 1661-1672. https://doi.org/10.1111/1365-2745.12316
Stark, S., Männistö, M. & Eskelinen, A. 2014. Nutrient availability and pH jointly constrain microbial extracellular enzyme activities in nutrient-poor tundra soils. Plant and Soil 383: 373 – 385. https://doi.org/10.1007/s11104-014-2181-y
Eskelinen, A. & Harrison, S. 2014. Exotic plant invasions under enhanced rainfall are constrained by soil nutrients and competition. Ecology 95: 682-692. https://doi.org/10.1890/13-0288.1
2013 —————————————————————————————————————————-
Kaarlejärvi, E., Eskelinen, A. & Olofsson, J. 2013. Herbivory prevents lowland plants benefiting from warmer and more fertile conditions at high altitudes. Functional Ecology 27: 1244–1253. https://doi.org/10.1111/1365-2435.12113
2012 —————————————————————————————————————————-
Eskelinen, A., Harrison, S. & Tuomi, M. 2012. Plant traits mediate consumer and nutrient control on plant community productivity and diversity. Ecology 93: 2705–2718. https://doi.org/10.1890/12-0393.1
Stark, S., Eskelinen, A. & Männistö, M. 2012. Regulation of microbial function and communities by soil pH, nitrogen availability, and herbivory in tundra ecosystems. Ecosystems 15: 18-33. https://doi.org/10.1007/s10021-011-9491-1
2011 —————————————————————————————————————————-
Ruotsalainen, A.L. & Eskelinen, A. 2011. Root fungal symbionts interact with above-ground mammalian herbivory, soil nutrient availability, and specific habitat conditions. Oecologia 166: 807-817. https://doi.org/10.1007/s00442-011-1928-x
2010 & Before —————————————————————————————————————–
Eskelinen, A. 2010. Resident functional composition mediates the impacts of nutrient enrichment and neighbour removal on plant immigration rates. Journal of Ecology 98: 540 – 550. https://doi.org/10.1111/j.1365-2745.2010.01654.x
Eskelinen, A., Stark, S. & Männistö, M. 2009. Links between plant community composition, soil organic matter quality and microbial communities in contrasting tundra habitats. Oecologia 161: 113 – 123. https://doi.org/10.1007/s00442-009-1362-5
Eskelinen, A. 2008. Herbivore and neighbour effects on tundra plants depend on species identity, nutrient availability and local environmental conditions. Journal of Ecology 96: 155 – 165. https://doi.org/10.1111/j.1365-2745.2007.01322.x
Eskelinen, A. and Oksanen, J. 2006. Changes in the abundance, composition and species richness of mountain vegetation in relation to summer grazing by reindeer. Journal of Vegetation Science 17: 245 – 254. https://doi.org/10.1111/j.1654-1103.2006.tb02443.x
Eskelinen. A. & Virtanen, R. 2005. Local and regional processes in low-productive mountain plant communities: the roles of seed and microsite limitation in relation to grazing. Oikos 110: 360 – 368. https://doi.org/10.1111/j.0030-1299.2005.13579.x
Virtanen, R., Eskelinen, A. & Gaare, E. 2003. Historical changes in species composition, abundance and diversity of alpine plant communities in Norway and Finland. In: Nagy, L., Grabherr, G., Körner, C. & Thompson, D.B.A. (eds.). Alpine Biodiversity in Europe. Ecological Studies, Vol. 167. Springer Verlag, Berlin Heidelberg. pp. 411 – 422.
Eskelinen Research Group 