Metacommunity theory unites community and spatial ecology. Recent innovations in sequencing technology may now allow us, for the first time, to confront this theory with entire trophic community data at large scales. We metabarcoded vertebrate eDNA from 320 ponds in England. Using this ‘novel community data’, we calculated the ‘internal structure’ of this metacommunity, which is the contributions of environmental, biotic, and spatial processes, as well as stochasticity, to the distribution of each species and the composition of each site. Our results suggest that community assembly processes in this pondscape do not conform to a single metacommunity paradigm, but rather vary among species and over space. Combining this variation with external knowledge of species biologies allows us to better understand the differential influences of environmental filtering, species interactions, and dispersal limitation on several of the vertebrate species, paving the way for the routine empirical study of metacommunities.