26Stable isotope and fatty acid analyses were used to study carbon sources for animals in a 27 submerged plant bed. Epiphytes growing on Potamogeton perfoliatus, sand microflora, and 28 alder leaves were the most important carbon sources. The most abundant macrophyte, P. 29 perfoliatus was unimportant as a food source. Modelling (IsoSource) showed that epiphytes 30were the most important food source for the most abundant benthic invertebrates, the isopod 31Asellus aquaticus (annual mean contribution 64%), the amphipod Gammarus pulex (66%), 32 and the gastropod Potamopyrgus antipodarum (83%). The mean annual contributions of sand 33 microflora were respectively 21, 19, and 9% and of alder leaves, 15, 15, and 8% for these 34 three species. The relative importance of carbon sources varied seasonally. The relative 35 contribution of epiphytes was lowest for all three grazer species in July: A. aquaticus 38%, G. 36 pulex 43%, and P. antipodarum 42%. . A decline in epiphyte biomass in summer may have 37 caused this switch to less attractive food sources. P. perfoliatus provided habitat and shelter 38 for consumers, but food was mainly supplied indirectly by providing space for attached 39 epiphytes, which are fast-growing and provide a highly nutritious food source.