Manipulating ungulate herbivory in temperate and boreal forests: effects on vegetation and invertebrates. A systematic review

Background: Livestock grazing and 'overabundance' of large wild herbivores in forested areas have long been perceived as conflicting with the aims of both silviculture and forest conservation; however, certain kinds of herbivory can help to maintain habitat values in forest ecosystems. Management of mammalian herbivory in protected forests can, therefore, be a critical tool for biodiversity conservation. The primary aim of this systematic review was to examine how forest vegetation and invertebrates are affected by manipulation of the grazing/browsing pressure by livestock or wild ungulates. The ultimate purpose was to investigate whether such manipulation is useful for conserving or restoring biodiversity in forest set-asides. Methods: We considered studies of manipulated ungulate herbivory in forests anywhere within the boreal and temperate zones, not only in protected areas but also in production forest. Non-intervention or alternative levels of intervention were used as comparators. Relevant outcomes included abundance, diversity and composition of plants and invertebrates, tree regeneration, and performance of focal/target species. Studies were mainly selected from a recent systematic map of the evidence on biodiversity effects of forest management relevant to protected areas. Additional studies were identified through updated searches online and in bibliographies of existing reviews. Relevant studies were critically appraised, and studies with low or unclear validity were excluded from the review. Quantitative outcomes were extracted from 103 articles, and summary effect sizes were derived by meta-analysis. Results: Most of the 144 studies included in the review had been conducted in North America, Europe or Australia/New Zealand. The intervention most commonly studied was experimental exclusion (or enclosure) of wild and/or domestic ungulates by fencing. Other studies examined culling of wild ungulates or compared forests long grazed by livestock to ungrazed forests. Effects on vegetation and invertebrates were reported in 135 and 23 of the studies, respectively. We found negative responses to herbivory in the abundance of understorey vegetation as a whole, woody understorey and bryophytes, and also in the species richness of woody understorey vegetation, whereas the richness of forbs and bryophytes responded positively. Several effects depended on ungulate origins: Understorey abundance responded negatively to livestock and to ungulates introduced into the wild, but not to native ones. In contrast, understorey species richness responded positively to livestock but not to wild ungulates. The duration and intensity of herbivory had few significant effects on vegetation-exceptions included woody understorey abundance and richness, which decreased with increasing duration and intensity, respectively. Among invertebrates we found negative responses to herbivory in the abundance of lepidopterans and spiders, but no significant effects on species richness. Conclusions: Our review revealed a large body of high-validity experimental studies on impacts of ungulate herbivory in forests. This evidence confirmed that manipulation of such herbivory is often highly influential on tree regeneration and on the abundance, diversity and composition of understorey vegetation. Nevertheless, we also identified important knowledge gaps-we found few studies of boreal areas, long-term herbivory effects, impacts on bryophytes, lichens and invertebrates, and effects of manipulation less radical than total exclusion of ungulates.