From protege to nurse plant:Establishment of thorny shrubs in grazed temperate woodlands

QuestionThorny shrubs play keystone roles in grazed ecosystems by defending non-protected plants against herbivores, but their establishment in grazed ecosystems is poorly understood. Which factors control establishment of recruits of thorny nurse shrubs in grazed temperate woodlands?LocationAncient grazed temperate woodlands (52 degrees 32'N, 6 degrees 36'E), The Netherlands.MethodsWe surveyed biotic and abiotic factors for saplings of thorny nurse shrubs in plots with and without saplings. To disentangle these factors, we performed a transplantation experiment over two growing seasons with nurse shrub saplings (Prunus spinosa and Crataegus monogyna) planted in two dominant vegetation types - tall unpalatable swards and short grazed lawns - half of them protected from herbivory via exclosures.ResultsPlots with shrub saplings had taller surrounding vegetation, higher soil pH and higher soil moisture than plots without saplings. These plots predominantly contained unpalatable sward species, while plots without saplings mainly contained palatable lawn species. After transplantation, sapling survival was higher in exclosures than in the open, and higher in sward exclosures than in lawn exclosures. Sapling growth was higher in swards than in lawns, higher inside than outside exclosures, and higher for Prunus than Crataegus, while browsing on saplings was higher in lawns.ConclusionUnpalatable swards form essential establishment niches for thorny shrubs in grazed temperate woodlands: they protect against herbivores before thorns fully develop in saplings, and sapling growth is better due to improved micro-environmental conditions. Once established and thorny, shrub saplings grow out of the protective range of the swards and in turn facilitate tree seedlings, which are essential for long-term persistence of grazed temperate woodlands. This study shows that nurse plants may start as proteges before becoming facilitators for other plants in a later life stage. This may be common for nurse plants in various ecosystems. We argue that improved understanding of establishment of nurse plants and their constraining factors is crucial for effective conservation and restoration in various ecosystems.