Spatial and temporal variability of river periphyton below a hypereutrophic lake and a series of dams.

Klamath River is described as an "upside-down" river due to its origins from the hypereutrophic Upper Klamath Lake (UKL) and hydrology that is heavily regulated by upstream dams. Understanding the lake and reservoir effects on benthic communities in the river can inform important aspects of its water quality dynamics. Periphyton samples were collected in May-November from 2004, 2006-2013 at nine long-term monitoring sites along 306 river km below UKL and a series of dams (n=299). Cluster analysis of periphyton assemblages identified three statistically different periphyton groups (denoted Groups 1-3). Group 1 occurred primarily in the upstream reach for June-October and had a higher percentage of sestonic species, including the cyanobacteria Aphanizomenon flos-aquae and Microcystis aeruginosa, consistent with the presence of upstream reservoirs. Group 2 had the highest relative biomass of diatoms and lowest relative biomass of cyanobacteria. Sites in the lower reach of the Klamath River fell into Group 2 in May-June and transitioning into Group 3 for July-October. Group 3 was dominated by nitrogen (N)-fixing species, including three diatoms (Epithemia sorex, Epithemia turgida, and Rhopalodia gibba) with cyanobacterial endosymbionts and the cyanobacterium Calothrix sp. Periphyton assemblages were strongly associated with temporal variations in flow conditions (e.g., decreasing flow from spring to fall) and spatial gradients in nutrient concentrations (e.g., decreasing from upstream to downstream). The inverse longitudinal relationship between periphyton biomass and nutrients may be explained by the ability of benthic N-fixers (Group 3) to overcome N limitation. Overall results showed a strong inverse relationship between the relative biomass of N-fixers and nitrogen concentrations and flow. This long-term dataset provides valuable insight into Klamath River's seasonal and longitudinal patterns of benthic algal communities and associated environmental variables. Our findings can inform river management decisions such as reducing upstream nutrient loads, setting flow regimes, and potential dam removals.