Untangling the relationships between morphology and phylogeny is key to building a reliable taxonomy, but is especially challenging for protists, where the existence of cryptic or pseudocryptic species makes finding relevant discriminant traits difficult. Here we use Hyalosphenia papilio (a testate amoeba) as a model species to investigate the contribution of phylogeny and phenotypic plasticity in its morphology. We study the response of H. papilio morphology (shape and pores number) to environmental variables in (i) a manipulative experiment with controlled conditions (water level), (ii) an observational study of a within-site natural ecological gradient (water level), and (iii) an observational study across 37 European peatlands (climate). We showed that H. papilio morphology is correlated to environmental conditions (climate and water depth) as well as geography, while no relationship between morphology and phylogeny was brought to light. The relative contribution of genetic inheritance and phenotypic plasticity in shaping morphology varies depending on the taxonomic group and the trait under consideration. Thus, our data call for a reassessment of taxonomy based on morphology alone. This clearly calls for a substantial increase in taxonomic research on these globally still under-studied organisms leading to a reassessment of estimates of global microbial eukaryotic diversity.

Ecology and Biodiversity Group Faculty of Science Utrecht University Padualaan 8 Utrecht 3585 CH The Netherlands Ecole Polytechnique Federale de Lausanne Ecological Systems Laboratory Lausanne CH 1015 Switzerland WSL Swiss Federal Institute for Forest Snow and Landscape Research Site Lausanne Lausanne CH 1015 Switzerland

Environment University of York York YO105DD United Kingdom Department of Zoology and Ecology Penza State University Krasnaya street 40 440026 Penza Russia

Laboratory of Evolutionary Protistology Institute of Biosciences University of Sao Paulo Matao Travessa 14 Cidade Universitaria Sao Paulo 05508 090 SP Brazil Biology Center Czech Academy of Sciences Branišovská 1160 31 37005 České Budějovice Czech Republic

Laboratory of Soil Biodiversity University of Neuchatel Rue Emile Argand 11 Neuchatel 2000 Switzerland

Laboratory of Soil Biodiversity University of Neuchatel Rue Emile Argand 11 Neuchatel 2000 Switzerland EPEP UMR 7144 CNRS UPMC Roscoff Biological Station Place Georges Teissier 29680 Roscoff France

Laboratory of Soil Biodiversity University of Neuchatel Rue Emile Argand 11 Neuchatel 2000 Switzerland Jardin Botanique de Neuchatel Chemin du Perthuis du Sault 58 Neuchatel 2000 Switzerland

Laboratory of Wetland Ecology and Monitoring and Department of Biogeography and Paleoecology Adam Mickiewicz University Krygowskiego 10 Poznan 61 680 Poland

Laboratory of Wetland Ecology and Monitoring and Department of Biogeography and Paleoecology Adam Mickiewicz University Krygowskiego 10 Poznan 61 680 Poland Institute of Plant Sciences and Oeschger Centre for Climate Change Research University of Bern Altenbergrain 21 CH 3013 Bern Switzerland

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