Bloom succession and nitrogen dynamics during snowmelt in a mid-order montane river

The Upper Clark Fork River (UCFR), Montana, a mid-order well-lit system with contemporary anthropogenic nitrogen (N) enrichment and natural geogenic sources of phosphorus (P), experiences annual algal blooms that influence ecosystem structure and function. This study was designed to assess the occurrence of riverine algal blooms (RABs) in the UCFR by characterizing the succession of periphyton and biogeochemical conditions following annual snowmelt runoff through autumnal baseflow conditions, and to provide a framework for assessing RAB progression in montane mid-order rivers more broadly. Using a 21-year database (2000–2020) collected over the growing season at three sites, historical assessment of the persistent and recurrent character of RABs in the UCFR showed that the magnitude of the summer bloom was, in part, moderated by snowmelt disturbance. Abundance and growth forms of benthic algae, along with river physicochemistry (e.g., temperature) and water chemistry (N and P concentration), were measured over the course of snowmelt recession for three years (2018–2020) at the same three sites. Results documented the onset of major blooms of the filamentous green algae Cladophora across all sites, commensurate with declines in dissolved inorganic N. Atomic N:P ratios of river water suggest successional transitions from P- to N-limitation associated with mid-season senescence of Cladophora and development of a secondary bloom of N-fixing cyanobacteria, dominated by Nostoc cf. pruniforme . Rates of N-fixation, addressed at one of the sites during the 2020 snowmelt recession, increased upon Cladophora senescence to a maximal value among the highest reported for lotic systems (5.80 mg N/m 2 /h) before decreasing again to background levels at the end of the growing season. Based on these data, a heuristic model for mid-order rivers responding to snowmelt disturbance suggests progression from phases of physical stress (snowmelt) to optimal growth conditions, to conditions of biotic stress later in the growing season. Optimal growth is observed as green algal blooms that form shortly after peak snowmelt, then transition to stages dominated by cyanobacteria and autochthonous N production later in the growing season. Accordingly, interactions among algal composition, reactive N abundance, and autochthonous N production, suggest successional progression from reliance on external nutrient sources to increased importance of autochthony, including N-fixation that susta