Vegetation patch dynamics in rangelands : how feedbacks between large herbivores, vegetation and soil fauna alter patches over space and through time

Abstract

AIM : Large herbivore grazing is a popular conservation management tool to promote vegetation structural diversity of rangelands. However, vegetation patch dynamics, that is, how patches of grazing-defended tall vegetation and grazer-preferred short lawns shift over space and time, is poorly understood. Here, we describe a new conceptual framework for patch dynamics within rangelands, combining theories of classical cyclical succession, self-organization and multitrophic feedbacks between grazers, vegetation and bioturbating soil fauna. LOCATION : We use the cattle-grazed salt marsh of the island Schiermonnikoog, The Netherlands, as a model system. The grazed salt marsh is characterized by distinct tall vegetation patches dominated by the grazing-defended rush Juncus maritimus and grazing-intolerant grass Elytrigia atherica, surrounded by a matrix of grazing lawn (dominated by Festuca rubra). THE FRAMEWORK : Based on previous observational and experimental studies, we propose a cyclical patch dynamic where plant species composition and structure transitions through four phases: patch initiation (a) occurs when the grazing-defended rush J. maritimus establishes in the grazed lawn. Patch establishment (b) follows when the grazing-intolerant grass E. atherica establishes in the patch due to associational defence by J. maritimus and produces a large amount of litter that attracts the key bioturbating amphipod Orchestia gammarellus. Patch expansion (c) occurs when O. gammarellus activities improve soil properties of the patch, which favours E. atherica growth, leading to E. atherica competitively displacing J. maritimus in the centre of the patch. Patch degeneration (d) follows when cattle enter the enlarged patch to consume E. atherica in the centre, trample the soil, displace O. gammarellus and decrease vegetation cover, opening space for grazing-lawn species to invade. The cycle restarts when remnants of the rush J. maritimus in the degenerated patches (or individuals recently established from seed dispersal) initiate new patches in the grazing lawn. SYNTHESIS : Our proposed patch-dynamic model provides a means to describe the mechanisms driving vegetation patch dynamics and serves as a foundation for further experimental and observational exploration, not only for this specific system, but more generally for grazed systems worldwide that show patches of typical grazing-defended and grazer-preferred vegetation.