Higher-order interactions mitigate direct negative effects on population dynamics of herbaceous plants during succession

Abstract

Plant succession is regulated by a combination of abiotic and biotic factors. However, previous studies of biotic drivers have focused overwhelmingly on direct pairwise species interactions, ignoring the likely prevalent higher-order interactions (HOIs) in natural systems. Climate also plays a significant role in determining successional dynamics with both direct effects and indirect effects via altered biotic interactions. Here we explored the relative effects of direct species interactions, HOIs, climate, and their interactions on population dynamics of herbaceous plants during 50 years of post-agricultural secondary succession and tested whether the inclusion of HOIs and climate data improved forecasts of population dynamics. Direct intraspecific interactions were competitive and prevalent across the 90 herbaceous plants examined, while direct interspecific interactions only affected populations of 29% species. HOIs, mainly arose from intraspecific HOIs of conspecifics, were mostly positive and thus largely mitigated the competitive effects of direct intraspecific interactions. Species with lower peak cover experienced stronger intraspecific competition and positive intraspecific HOIs of conspecifics. Direct interspecific interactions had neutral or facilitative effects on species with lower peak cover, and tended to have competitive effects on species with higher peak cover. Climate simultaneously influenced population dynamics both directly and indirectly via altered species interactions. Forecast performance was significantly improved with the inclusion of HOIs or climate for about half and one-third of species, respectively. Our study emphasizes the importance of HOIs, which largely mitigated direct competitive effects on population dynamics of herbaceous plants during succession. Teasing apart HOIs from direct species interactions substantially refined our understanding of successional dynamics of herbaceous plants and improved the accuracy of forecasting population dynamics during succession in a changing world.