1. Global change, such as exotic invasions, dramatically affects ecosystem functioning. However, the mechanisms behind the impacts are often unclear and despite extensive experimental work, we know little about the importance of biodiversity loss as a component of global change effects in real-world ecosystems. 2. We disentangled several mechanisms by which an exotic invader affected ecosystem functioning in East African drylands in Kenya and Ethiopia. We used structural equation modelling to separate direct effects of a woody invader Prosopis juliflora on a range of ecosystem functions from indirect effects mediated through changes in biodiversity (plant species richness) and ecosystem functioning (herbaceous biomass). We analysed effects on ecosystem functions linked to soil biogeochemical cycling and transfer of energy between trophic levels. 3. We found that the mean size of individual indirect effects mediated by biodiversity and herbaceous biomass was about twice as large as the mean size of individual direct effects of Prosopis on ecosystem functions, showing that indirect effects are an important component of the invader's overall environmental impacts. Changes in both herbaceous biomass and biodiversity were approximately equally important as drivers of indirect effects, indicating that we need to expand our view of indirect effects to consider a wider range of mechanisms. Simple univariate models failed to capture some Prosopis invader effects because positive direct effects on soil stability and predatory invertebrates were counteracted by negative indirect effects on biodiversity or biomass loss. The majority of effects were similar in both study systems. Suggestions that woody invaders are able to | 2661 Journal of Ecology LINDERS Et aL.

Woody alien plant species have been deliberately introduced globally in many arid and semi-arid regions, as they can provide services and goods to the rural poor. However, some of these alien trees and shrubs have become invasive over time, with important impacts on biodiversity, ecosystem services, and human well-being. Prosopis was introduced in Baringo County, Kenya, in the 1980s, but since then, it has spread rapidly from the original plantations to new areas. To assess land-use and land-cover (LULC) changes and dynamics in Baringo, we used a combination of dry and wet season Landsat satellite data acquired over a seven-year time interval between 1988–2016, and performed a supervised Random Forest classification. For each time interval, we calculated the extent of Prosopis invasion, rates of spread, gains and losses of specific LULC classes, and the relative importance of Prosopis invasion on LULC changes. The overall accuracy and kappa coefficients of the LULC classifications ranged between 98.1–98.5% and 0.93–0.96, respectively. We found that Prosopis coverage increased from 882 ha in 1988 to 18,792 ha in 2016. The highest negative changes in LULC classes were found for grasslands (−6252 ha; −86%), irrigated cropland (−849 ha; −57%), Vachellia tortilis-dominated vegetation (−3602 ha; −42%), and rainfed cropland (−1432 ha; −37%). Prosopis invasion alone directly accounted for over 30% of these negative changes, suggesting that Prosopis invasion is a key driver of the observed LULC changes in Baringo County. Although the management of Prosopis by utilization has been promoted in Baringo for 10–15 years, the spread of Prosopis has not stopped or slowed down. This suggests that Prosopis management in Baringo and other invaded areas in East Africa needs to be based on a more integrated approach.