The complexity underlying invasiveness precludes the identification of invasive traits: A comparative study of invasive and non-invasive heterocarpic Atriplex congeners.

Heterocarpy enables species to effectively spread under unfavourable conditions by producing two or more types of fruit differing in ecological characteristics. Although it is frequent in annuals occupying disturbed habitats that are vulnerable to invasion, there is still a lack of congeneric studies addressing the importance of heterocarpy for species invasion success. We compared two pairs of heterocarpic Atriplex species, each of them comprising one invasive and one non-invasive non-native congener. In two common garden experiments, we (i) simulated the influence of different levels of nutrients and population density on plants grown from different types of fruits and examined several traits that are generally positively associated with invasion success, and (ii) grew plants in a replacement series experiment to evaluate resource partitioning between them and to compare their competitive ability. We found that specific functional traits or competitiveness of species cannot explain the invasiveness of Atriplex species, indicating that species invasiveness involves more complex interactions of traits that are important only in certain ecological contexts, i.e. in specific environmental conditions and only some habitats. Interestingly, species trait differences related to invasion success were found between plants growing from the ecologically most contrasting fruit types. We suggest that fruit types differing in ecological behaviour may be essential in the process of invasion or in the general spreading of heterocarpic species, as they either the maximize population growth (type C fruit) or enhance the chance of survival of new populations (type A fruit). Congeners offer the best available methodical framework for comparing traits among phylogenetically closely related invasive and non-invasive species. However, as indicated by our results, this approach is unlikely to reveal invasive traits because of the complexity underlying invasiveness.