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The effects of woody plant encroachment and wildfire on plant species richness and composition: Temporal changes in a forest–steppe mosaic

Woody plant encroachment and wildfire may both have major effects on species richness and composition, yet 43 studies that assess these two factors in combination are rare. We asked the following specific questions. (1) How 44 did juniper cover change over a decade in the study area? (2) What are the effects of juniper encroachment and 45 subsequent wildfire on plant species richness, and (3) on vegetation composition? We studied a juniper encroached grassland hit by a wildfire. We assessed changes in plant species richness and 52 composition in burnt and unburnt grassland and juniper plots for eleven years following the wildfire. Yearly 53 vegetation sampling was performed in permanent quadrats of 1 m 2 and 25 m 2 using visual cover estimation. 54 Temporal changes of juniper cover, and species richness were evaluated with generalized linear mixed-effects 55 models. Compositional changes were analysed by non-metric multidimensional scaling and fidelity analysis. At the beginning the the presence of individual juniper shrubs increased species richness, and did not alter species composition. However, microsites covered by junipers were extremely species poor. Juniper growth during the study caused sharp decline in species richness at the edge of juniper shrubs and a shift in species composition compared to grassland plots. Wildfire increased species richness both in grassland and juniper plots. It caused only transient compositional responses in grasslands, but converted juniper habitats back to grassland state.