Ecosystem management often aims to maintain a diversity of habitats to benefit a large number of species within a landscape. We studied the effects of wetland management by lowintensity cattle-grazing and late-summer burning on marsh vegetation and globally declining anuran amphibians (frogs and toads) in a previously homogeneous reedbed. Burning effectively removed old reed and increased the variability of reed cover and marsh vegetation by the next spring. However, reed grew back strong in areas burned 2 or 3 years before the study, indicating that fire rejuvenates reedbeds. In contrast, cattle-grazing kept reed cover homogeneously low and created open water surfaces. The number of amphibian species and individuals decreased with mean reed cover and old reed density, and increased with variability in reed cover. Correspondingly, amphibian richness and counts were greatest in newly burned areas the next spring. In contrast, a year later, richness and counts were greatest in grazed-only areas, with large decreases in newly burned and control areas. Our results suggest that combined management with grazing and burning can create different habitat patches, some of which will be optimal for amphibians in one year, whereas other patches may become suitable in a subsequent year when successional changes alter previously optimal patches. To maximize optimal habitats, mosaic management should repeat burning once every 2 or 3 years in a rotational manner, and also maintain low-intensity cattle-grazing, which controls reeds and benefits amphibians more sustainably. Our study supports spatiotemporally varied management to facilitate habitat heterogeneity and complexity in dynamic landscapes.

Ecosystem restoration implies focusing on multiple trophic levels and ecosystem functioning, yet higher trophic levels, that is, animals, are less frequently targeted by restoration than plants. Habitat diversity, the spatial heterogeneity between and within habitat patches in a landscape, is a well-known driver of species diversity, and offers possible ways to increase species diversity at multiple trophic levels. We argue that habitat diversity is central in whole-ecosystem restoration as we review its importance, provide a practical definition for its components, and propose ways to target it in restoration. Restoration targeting habitat diversity is used commonly in aquatic ecosystems, mostly to increase the physical diversity of habitats, meant to provide more niches available to a higher number of animal species. To facilitate the uptake of habitat diversity in terrestrial ecosystem restoration, we distinguish between compositional and structural habitat diversity, because different animal groups will respond to different aspects of habitat diversity. We also propose four methods to increase habitat diversity: varying the starting conditions to obtain divergent successional pathways, emulating natural disturbances, establishing keystone structures, and applying ecosystem engineer species. We provide two case studies to illustrate how these components and methods can be incorporated in restoration. We conclude that targeting habitat diversity is a promising way to restore habitats for a multitude of species of animals and plants, and that it should become mainstream in restoration ecology and practice. We encourage the restoration community to consider compositional and structural habitat diversity and to specifically target habitat diversity in ecosystem restoration.

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