Abstract

Both earthworms and arbuscular mycorrhizal fungi (AMF) are important ecosystem engineers co-occurring in temperate grasslands. However, their combined impacts during grassland establishment are poorly understood and have never been studied. We used large mesocosms to study the effects of different functional groups of earthworms (i.e., vertically burrowing anecics vs. horizontally burrowing endogeics) and a mix of four AMF taxa on the establishment, diversity and productivity of plant communities after a simulated seed rain of 18 grassland species comprising grasses, non-leguminous forbs and legumes. Moreover, effects of earthworms and/or AMF on water infiltration and leaching of ammonium, nitrate and phosphate were determined after a simulated extreme rainfall event (40 l m−2). AMF colonisation of all three plant functional groups was altered by earthworms. Seedling emergence and diversity was reduced by anecic earthworms, however only when AMF were present. Plant density was decreased in AMF-free mesocosms when both anecic and endogeic earthworms were active; with AMF also anecics reduced plant density. Plant shoot and root biomass was only affected by earthworms in AMF-free mesocosms: shoot biomass increased due to the activity of either anecics or endogeics; root biomass increased only when anecics were active. Water infiltration increased when earthworms were present in the mesocosms but remained unaffected by AMF. Ammonium leaching was increased only when anecics or a mixed earthworm community was active but was unaffected by AMF; nitrate and phosphate leaching was neither affected by earthworms nor AMF. Ammonium leaching decreased with increasing plant density, nitrate leaching decreased with increasing plant diversity and density. In order to understand the underlying processes of these interactions further investigations possibly under field conditions using more diverse belowground communities are required. Nevertheless, this study demonstrates that belowground-aboveground linkages involving earthworms and AMF are important mediators of the diversity, structure and functioning of plant communities.

Highlights

  • In temperate grasslands, earthworms and arbuscular mycorrhizal fungi (AMF) are among the most important heterotrophic soil organisms by making up the dominant fraction of soil fauna [1] or forming symbiotic associations with the majority of land plants [2]

  • Earthworm communities in temperate grasslands in Europe usually comprise species belonging to three functional groups, anecics or vertical burrowers, endogeics or horizontal burrowers and epigeics or surface dwellers mainly distinguished because of their behaviour, activity zones and food preferences within the soil [14,15,16]

  • In temperate grasslands despite seeds present in the soil seed bank, a great amount of seedling recruitment occurs via seed rain which can amount to almost 11000 seeds m2 year21 [27]

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Summary

Introduction

Earthworms and arbuscular mycorrhizal fungi (AMF) are among the most important heterotrophic soil organisms by making up the dominant fraction of soil fauna [1] or forming symbiotic associations with the majority of land plants [2]. Because of their eminent influence on ecosystem characteristics and functions, both earthworms and AMF are considered as ecosystem engineers in many terrestrial ecosystems [3,4,5]. Through burrowing, casting and mixing of litter and soil, earthworms impact microbial activity and nutrient availability in the soil [28,29,30], modifying the soil structure by producing stable macropores and aggregates [31,32,33] and influence soil water characteristics by increasing water and nutrient infiltration in soils [34,35,36,37,38]

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