Natural variation of macrophyte vegetation of lowland streams at the regional level

In the present study, we present a synopsis of two macrophyte surveys of physiographic units in northwest Germany carried out over one decade. Data were used to test a set of hypotheses on macrophyte distribution at the regional level. Rank-frequency curves resembled the broken stick model. Twenty-one species of the 59 most frequent species occurred at high frequencies above 15%. Helophytes made up a high percentage (12 of 21) of the frequent species. Phalaris arundinacea was the most frequent species in both sampling periods. Most species showed no considerable change in frequency over time, among them the core hydrophytes. Spatial variation of species frequencies among physiographical units showed a unimodal distribution in relation to frequency. Spatial variation of frequencies of functional groups was significantly lower. Most uneven distribution among physiographical units was found in cryptogams. DCA ordinations of physiographical units showed a spatial gradient from alluvial plains to higher grounds units, which remained constant over time. CCA ordination of physiographical units in relation to environmental parameters identified two main axes, an altitudinal gradient and an alkalinity gradient. Species composition of units corresponded to the main landscape pattern of alluvial plains, glacial lowlands, and higher grounds on Mesozoic rock. Species diversity showed a complex behavior. Diverse units were found both in alluvial plains and glacial lowlands of intermediate elevation. The study may help defining regionally differentiated reference states for stream management, benchmarking indicator scores of species and avoiding application of assessment methods outside their range of applicability.