Notes: 1. We examined the contribution of algal cells to periphytic organic carbon and assessed the effects of variable biomass composition on the carbon : phosphorus (C : P) ratio of periphyton. We compiled more than 5000 published and unpublished observations of periphytic carbon : chlorophyll a (C : Chl) ratios, an index of algal prevalence, from a variety of substrata collected from lake and low-salinity coastal habitats. In addition, we converted estimates of algal biovolume into algal C to obtain an independent measure of cellular algal carbon in periphyton. This information was used in a model relating periphyton C : P ratio to algal cellular carbon, the algal C : P ratio, and the C : P ratio of non-algal organic matter in periphyton.2. The mean C : Chl ratio of periphyton (405) was relatively high with values in 〉25% of the samples exceeding 500. On average, 8.4% of total periphyton C was accounted for by C in algal cells. Only 15% of samples were found to have more than 15% periphyton C in cellular algal carbon. Our model showed a nonlinear relationship between periphytic C : P ratios and the C : P ratio of algal cells in the periphyton when non-algal organic matter was present. However, even at relatively low cellular algal C (〈10% of total C), algal C : P ratios can strongly affect the C : P ratio of periphyton as a whole (i.e. algal cells plus other organic matter).3. The high C : Chl ratios and the low biovolume-derived algal C of periphyton samples in our data set indicate that algal cells are typically a minor component of organic carbon in periphyton, However, this minor contribution would not preclude algal cellular stoichiometry from notably influencing periphyton C : P ratios.

Notes: 1. The aim of this study was to estimate patchiness in biomass and in the internal nutrient status of benthic algae on hard substrata (epilithon) in Lake Erken, Sweden, over different levels of distance, depth and time. Knowledge of the sources and scale of patchiness should enable more precise estimation of epilithic biomass and nutrient status for the entire lake. We focused on the horizontal scale, about which little is known.2. We sampled epilithon by SCUBA diving and used a hierarchical sampling design with different horizontal scales (cm, dm, 10 m, km) which were nested in two temporal scales (within and between seasons). We also compared two successive years and three sampling depths (0, 1 and 4 m). Biomass was measured as particulate carbon and chlorophyll a (Chl a) and internal nutrient status as carbon : nitrogen : phosphorus (C : N : P) ratios and as specific alkaline phosphatase activity (APA).3. Horizontal variation accounted for 60–80 and 7–70% of the total variation in biomass and in nutrient status, respectively, at all depths and during both years. Both small and large scales accounted for significant variation. We also found variation with time and depth. Biomass increased in autumn after a summer minimum, and the within-season variation was very high. The lowest biomass was found at 0 m depth. Both N and P limitation occurred, being higher in 1996 than in 1997 and decreased with depth.4. As a consequence, any sampling design must address variation with distance, depth and time when estimating biomass or nutrient limitation of benthic algae for an entire lake. Based on this analysis, we calculated an optimal sampling design for detecting change in the epilithic biomass of Lake Erken between different sampling days. It is important to repeat the sampling as often as possible, but also the large scales (10 m and km) and the dm scale should be replicated. Using our calculations as an example, and after a pilot study, an optimal sampling design can be computed for various objectives and for any lake.5. Short-term impact of the wind, light and nutrient limitation, and grazing, might be important in regulating the biomass and nutrient status of epilithic algae in Lake Erken. Patchiness in the nutrient status of algae was not coupled to the patchiness of biomass, indicating that internal nutrients and biomass were regulated by different factors.