Patterns in the Submerged Macrophyte Biomass of Lakes and the Importance of the Scale of Analysis in the Interpretation

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Abstract
The relative contributions of lake characteristics (i.e. alkalinity, chlorophyll A concentration, total phosphorus concentration, conductivity, and morphometry) and site characteristics (i.e. depth, littoral slope, exposure to waves, and underwater light levels) to the variability in submerged biomass were examined in 25 Canadian and American lakes. Lake-average submerged biomass is a function of water alkalinity and the lake-average littoral slope whereas site-specific biomass is a function of both site and lake characteristics. Plant biomass decreased with increasing slope and wave exposure and increased with increasing alkalinity and light levels. However, these relationships are complex because submerged biomass is also influenced by threshold phenomena (e.g. critical littoral slopes and transparency-dependent critical depths) that set limits to macrophyte colonization and because the relative contributions of the most relevant environmental factors studied (i.e. littoral slope, exposure, water transparency, and alkalinity) are depth dependent. By demonstrating the importance of lake-average and site-specific scales of variation and the existence of noncontinuous (e.g. threshold) regulation mechanisms the findings provide a new conceptual framework for the study of the relationship between submerged macrophytes, and their associated biota as well as their environment.