The relevance of bryophytes in the macrophyte-based reference conditions in Portuguese rivers

The identification of macrophyte reference communities represents a significant challenge in Mediterranean zones, where few aquatic taxa exist compared to other regions of the world. In the highly seasonal rivers of Portugal, we explored the contribution of bryophytes to enhance the compositional and ecological characterization of reference conditions. We performed partitioning, classification, and ordination analyses and obtained eight groups of bryophytic assemblages explained by water availability and quality. Groups A, B, and H assemblages are type-specific of the inland areas of the northern eco-region, reflecting the neutral minerotrophic currents with summer low currents. Groups C and D are type-specific of acidic oligotrophic shifting currents dependent of precipitation events in the mountainous streams of the North Atlantic eco-region. Groups E, F, and G are widespread in continental northern, central, and southern zones in neutral and highly mineralized watercourses with seasonal dryness. We propose bryophytes as a tool in the characterization of the reference conditions for two main reasons: (i) many of the recorded taxa are capable of withstanding the natural seasonal desiccation, and (ii) the type-specific bryophytes have perennial life-strategies and can be assessed at any time of year as a support for other type-specific macrophytes in compositional assessments of reference conditions.