Demographic and life-history variability across the range of a widespread herbthe role of environmental, geographical and genetic factors / Variabilidad demográfica y de historia vital en una planta de amplia distribución: el papel de los factores medioambientales, geográficos y genéticos

Résumé

Widespread species have traditionally received much less attention than rare and endemic ones. However, they are crucial in macroecological patterns and in ecosystem structure and functioning. Thus, understanding the characteristics that allow widespread organisms to extend over large areas has a high interest from both theoretical and applied perspectives. One of the most frequent hypotheses to explain the success of widespread plants is that they show much wider ecological niches, and thus a high life-history and demographic variability. However, studies are often very specific and carried out over small spatio-temporal scales, which hinders a general understanding of intraspecific variation in widespread taxa. In this thesis, we span a large spatio-temporal scale and a large environmental gradient to analyze the magnitude and the possible causes of natural variation in the in the range centre and the northern periphery of the widespread herb Plantago coronopus. More precisely, we analyze variability in population dynamics, life-history traits, and genetic diversity in up to 22 populations in Europe and North Africa. We aim to explore the relation of such variability with the position of populations within the species¿ range, since peripheral populations are traditionally expected to show a lower and more variable performance with respect to central populations. Additionally, we aim to analyze the effects of the most relevant environmental factors in population and individual performance at different spatial scales. In the first chapter, we found higher values in central populations in some vital rates, such as fecundity and growth, but recruitment and density were higher in northern peripheral populations, and there were no clear differences between regions in temporal variability of vital rates. Differences in population performance across the species¿ range seemed to be correlated with local precipitation and intraspecific competition. In the second chapter, differences in mean values and variability of vital rates between central and peripheral areas led to no differences in stochastic population growth rates. In addition, recruitment was the most influential vital rate for population growth rates at different spatial scales, and we found the same pattern of differentiation in population dynamics in response to environmental conditions within central and peripheral regions. In the third chapter, we reported high variation among populations in seed traits along a steep environmental stress gradient. Moreover, patterns in seed production were opposite at the fruit and the individual scale, as a strategy of populations to maximize fitness in each set of local conditions. Finally, in the fourth chapter, we found no relationship within populations between phenotypic variability and genetic diversity. Phenotypic variation was mainly shaped by precipitation variability, suggesting adaptive variation, whereas genetic diversity was correlated with the central vs. peripheral position, probably in close relation with some random demographic processes experienced by populations in the past. Despite genetic diversity was higher in central populations, our results contradicted classical hypotheses predicting a lower demographic performance towards species¿ range edges. In fact, environmental conditions seemed to have a higher influence on plant performance than the position of populations within the species¿ range, which calls for the necessity of distinguishing between geographical periphery and ecological marginality in demographic studies. Overall, our study highlights the versatility of P. coronopus at different spatial scales in response to varying environmental conditions, complementing similar findings of previous research on the same taxon at smaller spatial scales. Such life-history variability seems to be a key factor for widespread plants to extend over large and heterogeneous ranges.