Abstract

Understanding the factors underlying the co-occurrence of multiple species remains a challenge in ecology. Biotic interactions, environmental filtering and neutral processes are among the main mechanisms evoked to explain species co-occurrence. However, they are most often studied separately or even considered as mutually exclusive. This likely hampers a more global understanding of species assembly. Here, we investigate the general hypothesis that the structure of co-occurrence networks results from multiple assembly rules and its potential implications for grassland ecosystems. We surveyed orthopteran and plant communities in 48 permanent grasslands of the French Jura Mountains and gathered functional and phylogenetic data for all species. We constructed a network of plant and orthopteran species co-occurrences and verified whether its structure was modular or nested. We investigated the role of all species in the structure of the network (modularity and nestedness). We also investigated the assembly rules driving the structure of the plant-orthopteran co-occurrence network by using null models on species functional traits, phylogenetic relatedness and environmental conditions. We finally compared our results to abundance-based approaches. We found that the plant-orthopteran co-occurrence network had a modular organization. Community assembly rules differed among modules for plants while interactions with plants best explained the distribution of orthopterans into modules. Few species had a disproportionately high positive contribution to this modular organization and are likely to have a key importance to modulate future changes. The impact of agricultural practices was restricted to some modules (3 out of 5) suggesting that shifts in agricultural practices might not impact the entire plant-orthopteran co-occurrence network. These findings support our hypothesis that multiple assembly rules drive the modular structure of the plant-orthopteran network. This modular structure is likely to play a key role in the response of grassland ecosystems to future changes by limiting the impact of changes in agricultural practices such as intensification to some modules leaving species from other modules poorly impacted. The next step is to understand the importance of this modular structure for the long-term maintenance of grassland ecosystem structure and functions as well as to develop tools to integrate network structure into models to improve their capacity to predict future changes.

Highlights

  • Understanding the rules underlying species assembly is a key challenge in ecology (HilleRisLambers et al, 2012)

  • Orthopterans were present in module 1 (Chorthippus biguttulus, Mecostethus parapleurus, Stenobothrus lineatus and Stethophyma grossum) and 2 (Chrysochraon dispar, Euthystira brachyptera, Metrioptera roeselii and Omocestus viridulus) and in module 4 with Chorthippus albomarginatus

  • Other examples of modular co-occurrence networks were found among soil microbes (Barberan et al, 2012; Banerjee et al, 2015). Such organization was not encountered in plants and arbuscular mycorrhizal fungi (AMF) co-occurrence networks (Encinas-Viso et al, 2016). Contrary to this plant-AMF network where random encounters appear to drive species assembly, our results show that the interaction of multiple assembly rules can explain the modular organization of the plant-orthopteran co-occurrence network

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Summary

Introduction

Understanding the rules underlying species assembly is a key challenge in ecology (HilleRisLambers et al, 2012). A growing body of evidence suggests that different species assembly mechanisms can operate simultaneously and that they should be placed along a continuum (Amarasekare et al, 2004; Mouquet et al, 2005; Gravel et al, 2006; Leibold and McPeek, 2006; Fournier et al, 2016) Understanding this complexity of processes is key to better predict and manage species assemblages and their associated functions and services (Balvanera et al, 2006; Cadotte et al, 2011) and becomes increasingly critical in the current context of global change and biodiversity crisis (Koh et al, 2004). We explore to what extent the combination of species co-occurrence network and functional and phylogenetic approaches can provide new insights on how multiple rules interact to shape species assembly

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