“…In fact, it is well known that environmental factors are the main factors in establishing longitudinal distribution between interspecific and intraspecific lineages, including among closely related species (e.g., aquatic insects: Hildrew & Edington, 1979 ; Ogitani et al, 2011 ; Saito & Tojo, 2016 ; Saito et al, 2018 ; freshwater snails: Atkinson et al, 2012 ; freshwater fish: Morita et al, 2016 ). Using river systems as a target of study gives an advantage of easy quantitative evaluation of various environmental factors that are closely involved in differentiating the niches of organisms inhabiting rivers such as riverbed slope gradient, flow velocity, water depth, water quality, water temperature, riverbed substrate size, the abundance of organic matter and algae periphyton, and the degree of isolation (Okamoto et al, 2022 ). Under such circumstances, detailed longitudinal distribution patterns on the microhabitat scale of freshwater animals inhabiting rivers have been frequently discussed (Costa & Melo, 2008 ; Mauchart et al, 2017 ; Morita et al, 2016 ; Ogitani et al, 2011 ; Okamoto et al, 2022 ; Roberts & Angermeier, 2007 ; Saito & Tojo, 2016 ).…”