Abstract

Facilitation among plants mediated by grazers occurs when an unpalatable plant extends its protection against grazing to another plant. This type of indirect facilitation impacts species coexistence and ecosystem functioning in a large array of ecosystems worldwide. It has nonetheless generally been understudied so far in comparison with the role played by direct facilitation among plants. We aimed at providing original data on indirect facilitation at the community scale to determine the extent to which indirect facilitation mediated by grazers can shape plant communities. Such experimental data are expected to contribute to refining the conceptual framework on plant–plant–herbivore interactions in stressful environments. We set up a 2‐year grazing exclusion experiment in tropical alpine peatlands in Bolivia. Those ecosystems depend entirely on a few, structuring cushion‐forming plants (hereafter referred to as “nurse” species), in which associated plant communities develop. Fences have been set over two nurse species with different strategies to cope with grazing (direct vs. indirect defenses), which are expected to lead to different intensities of indirect facilitation for the associated communities. We collected functional traits which are known to vary according to grazing pressure (LDMC, leaf thickness, and maximum height), on both the nurse and their associated plant communities in grazed (and therefore indirect facilitation as well) and ungrazed conditions. We found that the effect of indirectly facilitated on the associated plant communities depended on the functional trait considered. Indirect facilitation decreased the effects of grazing on species relative abundance, mean LDMC, and the convergence of the maximum height distribution of the associated communities, but did not affect mean height or cover. The identity of the nurse species and grazing jointly affected the structure of the associated plant community through indirect facilitation. Our results together with the existing literature suggest that the “grazer–nurse–beneficiary” interaction module can be more complex than expected when evaluated in the field.

Highlights

  • Species interactions are known to drive species coexistence, their relative abundance, and ecosystem properties, such as productivity and resilience to perturbations (Agrawal et al, 2007)

  • We studied two nurse species with two different strategies to cope with grazing and we investigated the effect of indirect facilitation on the associated communities by manipulating the presence of herbivores

  • We showed that (1) the nurse species affects the capacity of the nurse to provide indirect facilitation to its associated communities and that (2) indirect facilitation can act at the community level by maintaining CWMLDMC and CWVHeight relatively constant between grazed and ungrazed conditions

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Summary

Introduction

Species interactions are known to drive species coexistence, their relative abundance, and ecosystem properties, such as productivity and resilience to perturbations (Agrawal et al, 2007). Positive interactions have been shown to play a central role in structuring plant communities (Brooker et al, 2008; Bruno, Stachowicz, & Bertness, 2003; Cavieres, Hernández-­ Fuentes, Sierra-A­ lmeida, & Kikvidze, 2016; Michalet et al, 2006), in maintaining ecosystem functions (Cardinale, Palmer, & Collins, 2002; Kéfi, Holmgren, & Scheffer, 2016; Kéfi, Rietkerk, van Baalen, & Loreau, 2007b), in promoting species richness (Gross, 2008) and biodiversity at the evolutionary scale (Valiente-­Banuet & Verdú, 2007) Determining their influence on the organization and dynamics of plant communities impacted by global changes is widely recognized as a topical challenge in plant science and ecology (Bulleri, Bruno, Silliman, & Stachowicz, 2016; Cavieres et al, 2014; Michalet, Schöb, Lortie, Brooker, & Callaway, 2014; Soliveres et al, 2015). Indirect facilitation stems from a reduction in a negative effect on the associated species caused by an intermediary species (Callaway, 2007), which can be a plant (Levine, 1999) or an animal, such as a domestic herbivore (Anthelme & Michalet, 2009)

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