Stream invertebrates use ‘drift’ or downstream transport in the water column to disperse, but they also actively move upstream against current. These movements have important role in re-colonization of empty stream reaches and in regulation of population size. For more than half a century, research of stream drift was stimulated by the problem of ‘drift paradox’, i.e. the question how upstream reaches of streams remain inhabited despite constant and often ample losses of individuals by drift. In this study we examined drift and upstream movements of the amphipod Gammarus fossarum through monthly sampling over one-year period at two sites in the upper and middle course of a small unaltered stream in the northwest Croatia. Differences in tendency to drift and to move upstream and effects of environmental factors (temperature, flow, population density and detritus dry weight) on the movement dynamics on both temporal and spatial scale were examined for three size categories (adults, subadults and juveniles). While seasonal variations of drift and upstream movements were similar at two examined sites, at the middle course we recorded higher density, and higher drift and upstream movement rates. In both drift and upstream movements, adults were the most represented size category. In subadults and juveniles, both type of movements depended on population density. Adult upstream movements peaked in November and May/June, with massive upstream movements in May in the middle course. Seasonality of diurnal and nocturnal drift was very similar at both sites, while seasonality of diurnal and nocturnal upstream movements differed between the sites. Out of four examined independent environmental factors, density and temperature had the most important positive effect on spatiotemporal variations of diurnal drift, while nocturnal drift was more strongly affected by density and flow. Temperature was the most significant driver of diurnal upstream movements, while nocturnal upstream movements were most significantly positively affected by density. Massive upstream movements were driven by overcrowding and enabled adults to colonize microhabitats where food is more abundant and competition is less severe.
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