The impact of non-reproductive plant species on assessments of community structure and species co-occurrence patterns

Aims: Studies of community structure and co-occurrence patterns rely on the premise that community data reflect where species successfully grow and which species they grow with. However, plant censuses generally do not distinguish between species with reproductive individuals and those only represented by non-reproductive individuals. We tested whether inclusion of non-reproductive species, which may not reflect success in that location, significantly impacts evaluations of community structure and co-occurrence. Location: Queen's University Biological Station, Ontario, Canada, old-field plant communities. Methods: We quantified the impact of non-reproductive species in two plant communities by comparing community structure and co-occurrence patterns when non-reproductive species were included or excluded. Results: Including non-reproductive species significantly increased plot-level species richness in both communities (54% and 13% increases), altered species evenness in both communities, significantly impacted beta-diversity among plots in one site, and disproportionately impacted assessments of diversity in species-rich plots. Excluding non-reproductive species resulted in reduced negative co-occurrence patterns in both communities, with a substantially larger impact in one community. In that community, the impact of non-reproductive species was even more pronounced when abundance data were used in analysis, and when pair-wise co-occurrence patterns were assessed. Additionally, including non-reproductive species drastically decreased the number of species pairs with perfect negative co-occurrence across sites, indicating that these species can add 'noise' to co-occurrence patterns. We examined possible explanations for the presence of non-reproductive species. In one community, non-reproductive species were 22 times less abundant (per plot) than reproductive species within plots, although they were not rare overall. Differences in the number of nonreproductive species per plot across our focal communities were not clearly driven by differences in clonality, stress from extreme weather or low N. While these patterns are consistent with the interpretation that non-reproductive species are present due to mass effects, this possibility requires further research. Conclusions: Including non-reproductive plant species in censuses can significantly impact assessments of community structure and species co-occurrence. The divergent impact of their inclusion on our