Abiotic and biotic factors structure species assembly in ecosystems both horizontally and vertically. However, the way community composition changes along comparable horizontal and vertical distances in complex three-dimensional habitats, and the factors driving these patterns, remains poorly understood. By sampling ant assemblages at comparable vertical and horizontal spatial scales in a tropical rainforest, we tested hypotheses that predicted differences in vertical and horizontal turnover explained by different drivers in vertical and horizontal space. These drivers included environmental filtering, such as microclimate (temperature, humidity, and photosynthetic photon flux density) and microhabitat connectivity (leaf area), which are structured differently across vertical and horizontal space. We found that both ant abundance and richness decreased significantly with increasing vertical height. Although the dissimilarity between ant assemblages increased with vertical distance, indicating a clear distance-decay pattern, the dissimilarity was higher horizontally where it appeared independent of distance. The pronounced horizontal and vertical structuring of ant assemblages across short distances is likely explained by a combination of microclimate and microhabitat connectivity. Our results demonstrate the importance of considering three-dimensional spatial variation in local assemblages and reveal how highly diverse communities can be supported by complex habitats.

Biology Centre of Czech Academy of Sciences Institute of Entomology Ceske Budejovice Czech Republic

Centre for Agri Environmental Research School of Agriculture Policy and Development University of Reading Reading UK

Department of Zoology University of Cambridge Cambridge UK

Institute of Culture and Environment Alaska Pacific University Anchorage Alaska USA

School of Biological and Behavioural Sciences Queen Mary University of London London UK

School of Ecology Sun Yat Sen University Guangzhou P R China

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