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Chironomids are keystone primary benthic consumers with semi‐aquatic life cycles. They support aquatic and terrestrial consumers at higher trophic levels by conveying dietary nutrients, such as fatty acids. In this study, we combined field sampling and laboratory experiments to examine the effects of environmental parameters, including diet, on fatty acid composition and metabolism in chironomid larvae and imagines. Results from 53 lakes showed that lake size, depth, dissolved organic carbon (DOC) concentrations, and trophic state had only marginal effects on the content of long‐chain polyunsaturated fatty acids (LC‐PUFA) in chironomids. Compound‐specific stable hydrogen isotope analyses confirmed that chironomids actively bioconvert dietary fatty acid precursors to LC‐PUFA in all lake types, independent of nutrient or DOC concentrations. Moreover, fatty acid‐specific stable carbon isotope data indicated that the diet of chironomids was subsidized, particularly in oligotrophic lakes in spring, by terrestrial C18 fatty acid precursors that were converted to LC‐PUFA. Data from feeding experiments further confirmed that decreased dietary availability of LC‐PUFA enhanced the conversion of dietary short‐chain precursors to LC‐PUFA. These results suggest that chironomids are PUFA regulators that can sustain LC‐PUFA levels under varying environmental conditions. Furthermore, our results indicate that they bioconvert terrestrial low‐quality material to high‐quality resources, which, via chironomid emergence, support terrestrial food webs. Chironomids are abundant and widespread, and thus, the trophic transfer of LC‐PUFA can have significant implications for the fitness and production of upper trophic level consumers in both aquatic and terrestrial ecosystems. [ABSTRACT FROM AUTHOR]