The complex structure of aquatic food webs emerges from a few assembly rules.

Food-web theory assumes that larger-bodied predators generally select larger prey. This allometric rule fails to explain a considerable fraction of trophic links in aquatic food webs. Here we show that food-web constraints result in guilds of predators that vary in size but have specialized on prey of the same size, and that the distribution of such specialist guilds explains about one-half of the food-web structure. We classified 517 pelagic species into five predator functional groups. Most of these follow three prey selection strategies: a guild following the allometric rule whereby larger predators eat larger prey and two guilds of specialists that prefer either smaller or larger prey than predicted by the allometric rule. Such coexistence of non-specialist and specialist guilds independent from taxa or body size points towards structural principles behind ecological complexity. We show that the pattern describes >90% of observed linkages in 218 food webs in 18 aquatic ecosystems worldwide. The pattern can be linked to eco-evolutionary constraints to prey exploitation and provides a blueprint for more effective food-web models.