Summary1. Global warming is predicted to dramatically alter communities' composition through differential colonization abilities, such as between sessile plants and their mobile herbivores. Novel interactions between previously non-overlapping species may, however, also be mediated by altered plants' responses to herbivore attack. 2. Syndromes of plant defences and tolerance are driven by inherited functional traits, biotic and abiotic conditions, and the geographical and historical contingencies affecting the community. Therefore, understanding climate change-driven herbivore responses and evolution towards a particular plant defence syndrome is key to forecasting species interactions in the near future. 3. In this paper, we first document variations in herbivory, and plant defences along altitudinal gradients that act as 'natural experiments'. We then use an empirical model to predict how specialist herbivore abundance may shift with respect to elevation in the near future. 4. Our field surveys and field experiment showed a decrease in herbivory with elevation. However, contrary to expectations, our meta-regression analyses showed that plant defences, particularly leaf toughness and flavonoid compounds, tend to be higher at high elevations, while secondary metabolites showed no clear trend with elevation. 5. Based on those results, we discuss how plant communities and species-specific plant defence syndromes will change in response to the climate-driven herbivore colonization of higher altitudes. Particularly, plant from high elevation, due to high protection against abiotic stress may be already ecologically fitted to resist the sudden increase in herbivory pressure that they will likely experience during global change.

Summary Symbioses between plants and fungi, fungi and ants, and ants and plants all play important roles in ecosystems. Symbioses involving all three partners appear to be rare. Here, we describe a novel tripartite symbiosis in which ants and a fungus inhabit domatia of an ant‐plant, and present evidence that such interactions are widespread. We investigated 139 individuals of the African ant‐plant Leonardoxa africana for occurrence of fungus. Behaviour of mutualist ants toward the fungus within domatia was observed using a video camera fitted with an endoscope. Fungi were identified by sequencing a fragment of their ribosomal DNA. Fungi were always present in domatia occupied by mutualist ants but never in domatia occupied by opportunistic or parasitic ants. Ants appear to favour the propagation, removal and maintenance of the fungus. Similar fungi were associated with other ant‐plants in Cameroon. All belong to the ascomycete order Chaetothyriales; those from L. africana formed a monophyletic clade. These new plant–ant–fungus associations seem to be specific, as demonstrated within Leonardoxa and as suggested by fungal phyletic identities. Such tripartite associations are widespread in African ant‐plants but have long been overlooked. Taking fungal partners into account will greatly enhance our understanding of symbiotic ant–plant mutualisms.

Understanding the functional economics that drives plant investment of resources requires investigating the interface between plant phenotypes and the variation in ecological conditions. While allocation to defence represents a large portion of the carbon budget, this axis is usually neglected in the study of plant economic spectrum. Using a novel geometrical approach, we analysed the co-variation in a comprehensive set of functional traits related to plant growth strategies, as well as chemical defences against herbivores on all 15 Cardamine species present in the Swiss Alps. By extracting geometrical information of the functional space, we observed clustering of plants into three main syndromes. Those different strategies of growth form and defence were also distributed within distinct elevational bands demonstrating an association between the functional space and the ecological conditions. We conclude that plant strategies converge into clear syndromes that trade off abiotic tolerance, growth and defence within each elevation zone.

The diversity of fungi along environmental gradients has been little explored in contrast to plants and animals. Consequently, environmental factors influencing the composition of fungal assemblages are poorly understood. The aim of this study was to determine whether the diversity and composition of leaf and root-associated fungal assemblages vary with elevation and to investigate potential explanatory variables. High-throughput sequencing of the Internal Transcribed Spacer 1 region was used to explore fungal assemblages along three elevation gradients, located in French mountainous regions. Beech forest was selected as a study system to minimise the host effect. The variation in species richness and specific composition was investigated for ascomycetes and basidiomycetes assemblages with a particular focus on root-associated ectomycorrhizal fungi. The richness of fungal communities associated with leaves or roots did not significantly relate to any of the tested environmental drivers, i.e. elevation, mean temperature, precipitation or edaphic variables such as soil pH or the ratio carbon∶nitrogen. Nevertheless, the ascomycete species richness peaked at mid-temperature, illustrating a mid-domain effect model. We found that leaf and root-associated fungal assemblages did not follow similar patterns of composition with elevation. While the composition of the leaf-associated fungal assemblage correlated primarily with the mean annual temperature, the composition of root-associated fungal assemblage was explained equally by soil pH and by temperature. The ectomycorrhizal composition was also related to these variables. Our results therefore suggest that above and below-ground fungal assemblages are not controlled by the same main environmental variables. This may be due to the larger amplitude of climatic variables in the tree foliage compared to the soil environment.

To cope with environmental challenges, plants produce a wide diversity of phytochemicals, which are also the source of numerous medicines. Despite decades of research in chemical ecology, we still lack an understanding of the organization of plant chemical diversity across species and ecosystems. To address this challenge, we hypothesized that molecular diversity is not only related to species diversity, but also constrained by trophic, climatic, and topographical factors. We screened the metabolome of 416 vascular plant species encompassing the entire alpine elevation range and four alpine bioclimatic regions in order to characterize their phytochemical diversity. We show that by coupling phylogenetic information, topographic, edaphic, and climatic variables, we predict phytochemical diversity, and its inherent composition, of plant communities throughout landscape. Spatial mapping of phytochemical diversity further revealed that plant assemblages found in low to midelevation habitats, with more alkaline soils, possessed greater phytochemical diversity, whereas alpine habitats possessed higher phytochemical endemism. Altogether, we present a general tool that can be used for predicting hotspots of phytochemical diversity in the landscape, independently of plant species taxonomic identity. Such an approach offers promising perspectives in both drug discovery programs and conservation efforts worldwide.

No abstract

In ant -plant symbioses, plants provide symbiotic ants with food and specialized nesting cavities (called domatia). In many ant -plant symbioses, a fungal patch grows within each domatium. The symbiotic nature of the fungal association has been shown in the ant-plant Leonardoxa africana and its protective mutualist ant Petalomyrmex phylax. To decipher trophic fluxes among the three partners, food enriched in 13 C and 15 N was given to the ants and tracked in the different parts of the symbiosis up to 660 days later. The plant received a small, but significant, amount of nitrogen from the ants. However, the ants fed more intensively the fungus. The pattern of isotope enrichment in the system indicated an ant behaviour that functions specifically to feed the fungus. After 660 days, the introduced nitrogen was still present in the system and homogeneously distributed among ant, plant and fungal compartments, indicating efficient recycling within the symbiosis. Another experiment showed that the plant surface absorbed nutrients (in the form of simple molecules) whether or not it is coated by fungus. Our study provides arguments for a mutualistic status of the fungal associate and a framework for investigating the previously unsuspected complexity of food webs in ant -plant mutualisms.

Nicole M. van Dam and Roberto Salguero-Gómez joint last authorship.