Ecosystems are increasingly prone to climate extremes, such as drought, with long-lasting effects on both plant and soil communities and, subsequently, on carbon (C) cycling. However, recent studies underlined the strong variability in ecosystem's response to droughts, raising the issue of nonlinear responses in plant and soil communities. The conundrum is what causes ecosystems to shift in response to drought. Here, we investigated the response of plant and soil fungi to drought of different intensities using a water table gradient in peatlands-a major C sink ecosystem. Using moving window structural equation models, we show that substantial changes in ecosystem respiration, plant and soil fungal communities occurred when the water level fell below a tipping point of -24 cm. As a corollary, ecosystem respiration was the greatest when graminoids and saprotrophic fungi became prevalent as a response to the extreme drought. Graminoids indirectly influenced fungal functional composition and soil enzyme activities through their direct effect on dissolved organic matter quality, while saprotrophic fungi directly influenced soil enzyme activities. In turn, increasing enzyme activities promoted ecosystem respiration. We show that functional transitions in ecosystem respiration critically depend on the degree of response of graminoids and saprotrophic fungi to drought. Our results represent a major advance in understanding the nonlinear nature of ecosystem properties to drought and pave the way towards a truly mechanistic understanding of the effects of drought on ecosystem processes.

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Mid‐ to high‐latitude peatlands are a major terrestrial carbon stock but become carbon sources during droughts, which are increasingly frequent as a result of climate warming. A critical question within this context is the sensitivity to drought of peatland microbial food webs. Microbiota drive key ecological and biogeochemical processes, but their response to drought is likely to impact these processes. Peatland food webs have, however, been little studied, especially the response of microbial predators. We studied the response of microbial predators (testate amoebae, ciliates, rotifers, and nematodes) living in Sphagnum moss carpet to droughts, and their influence on lower trophic levels and on related microbial enzyme activity. We assessed the impact of reduced water availability on microbial predators in two peatlands using experimental (Linje mire, Poland) and natural (Forbonnet mire, France) water level gradients, reflecting a sudden change in moisture regime (Linje), and a typically drier environment (Forbonnet). The sensitivity of different microbial groups to drought was size dependent; large sized microbiota such as testate amoebae declined most under dry conditions (−41% in Forbonnet and −80% in Linje). These shifts caused a decrease in the predator–prey mass ratio (PPMR). We related microbial enzymatic activity to PPMR; we found that a decrease in PPMR can have divergent effects on microbial enzymatic activity. In a community adapted to drier conditions, decreasing PPMR stimulated microbial enzyme activity, while in extreme drought experiment, it reduced microbial activity. These results suggest that microbial enzymatic activity resulting from food web structure is optimal only within a certain range of PPMR, and that different trophic mechanisms are involved in the response of peatlands to droughts. Our findings confirm the importance of large microbial consumers living at the surface of peatlands on the functioning of peatlands, and illustrate their value as early warning indicators of change.

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Please cite this article as: Swindles, G.T., Lamentowicz, M., Reczuga, M., Galloway, J.M.,Palaeoecology of testate amoebae in a tropical peatland, European Journal of Protistology (2015), http://dx.doi.org/10.1016/j. ejop.2015.10.002 This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. A c c e p t e d M a n u s c r i p t 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 2 Abstract 21We present the first detailed analysis of subfossil testate amoebae from a tropical 22 peatland. Testate amoebae were analysed from a 4-m peat core from western Amazonia 23 (Peru) and a transfer function developed from the site was applied to reconstruct changes in 24 water table over the past ca. 8,000 years. Testate amoebae were in very low abundance in the 25 core, especially in the lower 125 cm, due to a combination of poor preservation and 26 obscuration by other organic matter. A modified preparation method enabled at least 50 27 testate amoebae to be counted in each core sample. The most abundant taxa preserved include 28