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The study of biological invasions has long considered species invasiveness and community invasibility as separate questions. Only recently, there is an increasing recognition that integrating these two questions offers new insights into the mechanisms of biological invasions. This recognition has renewed the interest in two long-standing and seemingly contradictory hypotheses proposed by Darwin: phylogenetic relatedness of invaders to native communities is predicted to promote naturalization because of appropriate niche-adaptation but is at the same time predicted to hamper naturalization because of niche overlap with native species. The latter is known as Darwin's naturalization hypothesis. Global. We review the studies that have tested these hypotheses and summarize their largely inconsistent outcomes. We argue that most of the inconsistency arises from discrepancies in the applied conceptual frameworks and analytical approaches and not from different model organisms and different ecological contexts. First, observed patterns and results can be seriously flawed by different spatial and phylogenetic scales, which do not equally reveal community assembly mechanisms. Second, different studies have used different metrics, which may test for different specific hypotheses. Thus, we propose a set of metrics derived from the alpha niche concept to measure invaders relatedness to native communities. Finally, approximating species niche differentiation from phylogenetic relatedness is not exempt of assumptions, and invasive species naturalization may result from various ecological mechanisms of biotic resistance that are not necessarily revealed by species phylogeny alone. The quest for resolving the conundrum of Darwin's naturalization hypothesis will only be successful if appropriate scales, metrics and analytical tests are thoroughly considered. We give several recommendations and suggest, whenever possible, to use trait-based measurements of species dissimilarity as the most promising avenue to unravel the mechanisms driving alien species invasions.

If natural communities are assembled according to deterministic rules, coexisting species will represent a nonrandom subset of the potential species pool. We tested for signatures of assembly rules in the distribution of species' traits in Pacific rockfish (Sebastes spp.) assemblages. We used morphology, dietary niche (estimated with stable nitrogen isotopes), and distribution data to identify traits that relate to local-scale resource use (the α-niche) and to environmental gradients (the β-niche). We showed that gill raker morphology was related to trophic position (an α-niche axis), while relative eye size was associated with depth habitat (a β-niche axis). We therefore hypothesized that, within assemblages of coexisting rockfish species, the gill raker trait would be overdispersed (evenly spaced) due to limiting similarity, while relative eye size would be clustered due to environmental filtering. We examined the evolutionary relatedness of coexisting species to ask whether phylogenetic community structure and trait distributions gave similar indications about the roles of assembly processes. We tested the trait distributions and phylogenetic structure of 30 published rockfish assemblages against a null model of random community assembly. As predicted, the gill raker trait tended to be more evenly spaced than expected by chance, as did overall body size, while relative eye size was more clustered than expected. Phylogenetic community structure appeared to reflect historical dispersal and speciation and did not provide consistent support for assembly rules. Our results indicate that rockfish community assembly is nonrandom with regard to species' traits and show how distinguishing traits related to the α- and β-niches and incorporating functional morphology can provide for powerful tests of assembly rules.

Species assembly patterns of local communities are shaped by the balance between multiple abiotic/biotic filters and dispersal that both select individuals from species pools at the regional scale. Knowledge regarding functional assembly can provide insight into the relative importance of the deterministic and stochastic processes that shape species assembly. We evaluated the hierarchical roles of the a niche and β niches by analyzing the influence of environmental filtering relative to functional traits on geographical patterns of tree species assembly in mid-latitude forests. Using forest plot datasets, we examined the a niche traits (leaf and wood traits) and β niche properties (cold/drought tolerance) of tree species, and tested non-randomness (clustering/over-dispersion) of trait assembly based on null models that assumed two types of species pools related to biogeographical regions. For most plots, species assembly patterns fell within the range of random expectation. However, particularly for cold/drought tolerance-related β niche properties, deviation from randomness was frequently found; non-random clustering was predominant in higher latitudes with harsh climates. Our findings demonstrate that both randomness and non-randomness in trait assembly emerged as a result of the a and β niches, although we suggest the potential role of dispersal processes and/or species equalization through trait similarities in generating the prevalence of randomness. Clustering of β niche traits along latitudinal climatic gradients provides clear evidence of species sorting by filtering particular traits. Our results reveal that multiple filters through functional niches and stochastic processes jointly shape geographical patterns of species assembly across mid-latitude forests.