Freshwater reefs as mesohabitats for the assessment of diel invertebrate drift patterns

Freshwater reefs (known as tufa barriers) are special karst features recognized for highly heterogeneous habitat structures, complex hydrogeological features, and unique macrozoobenthos drift (downstream dispersal) patterns. Our study objective was to investigate diel and seasonal drift patterns between barriers and pools, both composed of moss-rich and fishless mesohabitat types, aligned on a small spatial scale within the karst, tufa-precipitating Plitvice Lakes hydrosystem. We monthly sampled drift at the two mesohabitat types (barriers and pools) during midday and dusk and examined quantitative and qualitative drift compositions, including drifting invertebrates, moss, and associated particulate organic and inorganic matter (APOIM). Barriers displayed higher invertebrate drift densities than those of pools. The same pattern was observed for moss and APOIM. At both mesohabitat types, invertebrate drift showed peak but highly variable densities during late spring and summer (mean >100 individuals m-3), whereas during late winter and early spring the drift densities were 5-fold lower than those densities. The nonmetric multidimensional scaling analysis revealed that invertebrate drift seasonality was influenced by seasonal drift patterns of aquatic moss and moss-dwelling invertebrate taxa. Adult and/or larval Riolus spp. and larval Hemerodromia spp. were found to be the most significant for the separation of midday and dusk within the NMDS ordination of barriers and pools. At barriers, midday drift densities of invertebrates, moss, and APOIM were higher than the respective dusk records. Within pools, invertebrate drift was largely aperiodic. We suggest that increased midday and/or aperiodic drift are a consequence of the lack of fish between barrier- and pool-mesohabitats. Our results further indicated that aquatic invertebrates inhabiting fast-flowing barriers and slow-flowing pools mostly exhibit “passive drift” mediated by transport agents such as water flow and dislodged aquatic vegetation. The observed spatio-temporal drift patterns are also likely influenced by ontogenetic shifts in drift periodicity (i.e., shifts depending on the development stage and morphological characteristics of the individual taxa) as well as benthic distribution of moss-dwelling invertebrate taxa. We can conclude that biotic (vs. abiotic) controls of drift are likely minimized in the fishless case of the freshwater reefs and associated barrier–pool sequences within Plitvice Lakes hydrosystem.