Temperature seasonality and nutrient enrichment drive intra-annual community turnover in global grasslands

In many grasslands, species with specific traits occupy unique temporal positions within communities. Such intra-annual segregation is predicted to be greatest in systems with high intra-annual climate variability because fluctuating environmental conditions provide opportunities for temporal niche partitioning among species. However, because most studies on intra-annual community dynamics have been conducted at individual sites, relationships between intra-annual climate variability and seasonal community dynamics at global scales have not yet been identified. Furthermore, the same characteristics that promote species-specific responses to fluctuations in environmental conditions may also drive species-specific responses to global change drivers such as eutrophication. Research provides evidence that eutrophication alters inter-annual plant community dynamics yet understanding of how it alters intra-annual dynamics remains limited. We used early-season and late-season compositional data collected from 10 grassland sites around the world to ask how intra-annual variability in precipitation and temperature as well as nutrient enrichment shape intra-annual species segregation, or seasonal {beta}-diversity, in plant communities. We also assessed whether changes in the abundances of specific functional groups including annual forbs, perennial forbs, C3 and C4 graminoids, and legumes underpin compositional differences between early- and late-season communities and treatments. We found that intra-annual temperature variability and seasonal {beta}-diversity were positively related but observed no relationship between intra-annual precipitation variability and seasonal {beta}-diversity. This suggests that positive relationships between -diversity and intra-annual temperature variability identified in earlier studies may be underpinned by the positive influence of intra-annual temperature variability on temporal segregation of species within growing seasons. We found that nutrient enrichment increased seasonal {beta}-diversity via increased turnover of species between early- and late-season communities. This finding mirrors patterns observed at inter-annual scales and suggests fertilization can alter compositional dynamics via similar mechanisms at varied temporal scales. Finally, fertilization reduced the abundance of C4 graminoids and legumes and eliminated intra-annual differences in these groups. In contrast, fertilization resulted in intra-annual differences in C3 graminoids which were not observed in control conditions, and increased abundance of C3 graminoids and annual forbs overall. Our study provides new insight into how intra-annual climate variability and nutrient enrichment influence biodiversity and seasonal dynamics in global grasslands.