Potential drivers of changing ecological conditions in English and Welsh rivers since 1990.

River invertebrate communities across Europe have been changing in response to variations in water quality over recent decades, but the underlying drivers are difficult to identify because of the complex stressors and environmental heterogeneity involved. Here, using data from ∼4000 locations across England and Wales, collected over 29 years, we use three approaches to help resolve the drivers of spatiotemporal variation in the face of this complexity: i) mapping changes in invertebrate richness and community composition; ii) structural equation modelling (SEM) to distinguish land cover, water quality and climatic influences; and iii) geographically weighted regression (GWR) to identify how the apparent relationships between invertebrate communities and abiotic variables change across the area. Mapping confirmed widespread increases in richness and the proportion of pollution-sensitive taxa across much of England and Wales. It also revealed regions where pollution-sensitive taxa or overall richness declined, the former primarily in the uplands. SEMs confirmed strong increases in average biochemical oxygen demand and nutrient concentrations related to urban and agricultural land cover, but only a minority of land cover's effect upon invertebrate communities was explained by average water chemistry, highlighting potential factors such as episodic extremes or emerging contaminants. GWR identified strong geographical variation in estimated relationships between macroinvertebrate communities and environmental variables, with evidence that the estimated negative impacts of nutrients and water temperature were increasing through time. Overall the results are consistent with widespread biological recovery of Britain's rivers from past gross organic pollution, whilst highlighting declines in some of the most diverse and least impacted streams. Modelling points to a complex and changing set of drivers, highlighting the multifaceted impacts of catchment land cover and the evolving role of different stressors, with the relationship to gross organic pollution weakening, whilst estimated nutrient and warming effects strengthened.