The relatively poor simulation of the below-ground processes is a severe drawback for many ecosystem models, especially when predicting responses to climate change and management. For a meaningful estimation of ecosystem production and the cycling of water, energy, nutrients and carbon, the integration of soil processes and the exchanges at the surface is crucial. It is increasingly recognized that soil biota play an important role in soil organic carbon and nutrient cycling, shaping soil structure and hydrological properties through their activity, and in water and nutrient uptake by plants through mycorrhizal processes. In this article, we review the main soil biological actors (microbiota, fauna and roots) and their effects on soil functioning. We review to what extent they have been included in soil models and propose which of them could be included in ecosystem models. We show that the model representation of the soil food web, the impact of soil ecosystem engineers on soil structure and the related effects on hydrology and soil organic matter (SOM) stabilization are key issues in improving ecosystem-scale soil representation in models. Finally, we describe a new core model concept (KEYLINK) that integrates insights from SOM models, structural models and food web models to simulate the living soil at an ecosystem scale.

A N Severtsov Institute of Ecology and Evolution RAS Moscow Russia

Agrosphere Institute IBG Forschungszentrum Jülich GmbH Jülich Germany

BC3 Basque Centre for Climate Change Scientific Campus of the University of the Basque Country Bilbao Bizkaia Spain

Biogeochemistry Group Forest Soils and Biogeochemistry Swiss Federal Research Institute WSL Birmensdorf Switzerland

Biogeography and Global Change National Museum of Natural Sciences Spanish National Research Council Madrid Spain

Comparative Plant and Fungal Biology Royal Botanic Gardens Kew London UK

CREAF Cerdanyola del Vallès Spain

Departamento de Ecología y Biología Animal Universidad de Vigo Vigo Spain

Department of Biodiversity Conservation and Ecosystem Restoration ARAID IPE CSIC Jaca Spain

Department of Biogeochemistry and Soil Quality Norwegian Institute of Bioeconomy Research Aas Norway

Department of Biology Plants and Ecosystems Universiteit Antwerpen Antwerpen Belgium

Department of Bioscience Aarhus University Silkeborg Denmark

Department of Ecological Science Vrije Universiteit Amsterdam Amsterdam Netherlands

Department of Forest Ecology and Management Swedish University of Agricultural Sciences Umeå Sweden

Earth and Life Institute UCLouvain Louvain la Neuve Belgium

Forest Research Alice Holt Lodge Farnham UK

Forest Soils and Biogeochemistry Swiss Federal Research Institute WSL Birmensdorf Switzerland

Groningen Institute for Evolutionary Life Sciences University of Groningen Groningen Netherlands

IKERBASQUE Basque Foundation for Science Bilbao Spain

Institute for Crop and Soil Science Julius Kühn Institut Braunschwei Germany

Institute for Environmental Studies Charles University Prague Czech Republic

Institute of Ecology and Earth Sciences University of Tartu Tartu Estonia

Institute of Ecology and Environmental Sciences IRD UPEC CNRS INRA Sorbonne Université Paris France

Institute of Forest Ecology Department of Forest and Soil Sciences University of Natural Resources and Life Sciences Vienna Austria

International Institute for Applied Systems Analysis IIASA Laxenburg Austria

OVAM Flemish Institute for Materials and Soils Mechelen Belgium

Slovenian Forestry Institute Ljubljana Slovenia

Universitat Autònoma de Barcelona Cerdanyola del Vallès Spain

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