Whole genome duplication increases ecological niche breadth of the perennial herb Urtica dioica

Environmental niche modelling and associated species distribution models often ignore intraspecific ecological differentiation linked with ploidal differentiation, i.e. the occurrence of several cytotypes of the same species. Here we tested the hypotheses that (i) the tetraploid (4x) cytotype is characterized by a broader ecological niche than the diploid (2x), and (ii) niche differentiation at the regional scale can serve as a proxy measure for differentiation at a smaller landscape scale in Urtica dioica, a perennial herbaceous plant with two prevailing cytotypes (2x and 4x). We focused on southern Moravia (Czech Republic), an area with a high diversity of vegetation types and the occurrence of both diploid and tetraploid cytotypes of U. dioica. To explore the geographical distribution, ecological preferences and habitat affinities of the diploid and tetraploid plants, we collected individuals at two different spatial scales: regional (similar to 1200 km(2)) and landscape (9.1 km(2)). We used flow cytometry to determine the ploidy level of all the plants collected and determined the distribution of the different cytotypes both in geographical and ecological space. We also compared the habitat affinities of the diploid and tetraploid plants. Analyses of ploidy revealed 93 diploids and 325 tetraploids (regional scale: 42 diploids, 85 tetraploids; landscape scale: 51 diploids and 240 tetraploids). Diploid plants occurred only in the wetter parts of the landscape and this pattern was very similar at both scales. In contrast, tetraploids occurred along the whole moisture gradient at both scales. Diploids and tetraploids differed also in their habitat affinities. While diploids occurred relatively more frequently in alluvial and wet forests, managed broadleaved forests, alluvial and wet meadows, and wetlands and riverbanks, tetraploids preferred oak and oak-hornbeam forests, ravine forests, actively managed meadows, human settlements and vineyards. Our data clearly show an ecological niche differentiation between diploid and tetraploid cytotypes of Urtica dioica. While tetraploids have a broad ecological tolerance and consequently a wide geographical distribution, diploids occur only in a narrow range of ecological conditions and their distribution is restricted to areas with a shallow water table in the bottoms of valleys. Thus, ploidal differentiation, confirmed for many plant species, should be included in environmental niche modelling and species distribution models.