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Size-structured food webs form integrated trophic systems where energy is being channelled from small to large consumers. Empirical evidence suggests that size structure prevails in aquatic ecosystems while in terrestrial food webs trophic level is largely independent of body size.Compartmentalisation of energy channeling according to size classes of consumers was suggested as a mechanism that underpins functioning and stability of terrestrial food webs including those belowground, but their structure has not been empirically assessed across the whole size spectrum.Here we used stable isotope analysis and metabolic regressions to describe size structure and energy use in eight belowground communities with consumers spanning 12 orders of magnitude in living body mass, from protists to earthworms. We showed a community-wide decline in trophic level with body mass in invertebrates and a remarkable non-linearity in community metabolism and trophic positions across all size classes. Specifically, we found that correlation between body mass and trophic level is positive in small-sized (protists, nematodes, arthropods below 1 µg in body mass), neutral in medium-sized (arthropods of 1 µg to 1 mg) and negative in large-sized consumers (large arthropods, earthworms), suggesting that these groups form compartments with different trophic organization. Based on this pattern, we propose a concept of belowground food webs being composed of (1) size-structured micro-food web driving fast energy channeling and nutrient release, e.g. in microbial loop, (2) arthropod macro-food web with no clear correlation between body size and trophic level, hosting soil arthropod diversity and subsidizing aboveground predators, and (3) 'trophic whales', sequestering energy in their large bodies and restricting its propagation to higher trophic levels in belowground food webs. The three size compartments are based on a similar set of basal resources, but contribute to different ecosystem-level functions and respond differently to variations in climate, soil characteristics and land use. We suggest that widely used vision of resource-based energy channeling in belowground food webs can be complemented with size-based energy channeling, where ecosystem multifunctionality, biodiversity and stability is supported by a balance across individual size compartments.

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Trophic resource partitioning is one of the main drivers of adaptive radiation. The evolutionary diversification of large African barbs, the genus Labeobarbus, seems to be related to mouth polymorphism. The chisel-mouthed or scraping phenotype has repeatedly evolved within Labeobarbus. At least five ecomorphs with a scraping mouth morphology were detected in the waters of the Ethiopian Highlands and can be provisionally classified into two groups: (i) “Varicorhinus”-like, and (ii) “Smiling”-like. Previously, all Labeobarbus with a scraping-mouth morphology were considered to be periphyton feeders. Using data on morphology, diet and stable isotope ratios (C and N), we addressed the question: does a scraping-mouth morphology predict feeding on periphyton? Our study revealed that five scraper ecomorphs exhibited three main feeding modes: (i) periphyton-eating, (ii) herbivory–detritivory, and (iii) insectivory. Two cases of the parallel divergence of sympatric ecomorphs with distinct feeding modes (herbivory–detritivory vs. insectivory) were revealed in two geographically isolated basins. A significant difference in δ15N values was detected among sympatric scraper ecomorphs. A periphytonophagous scraper was rich in δ15N values that are comparable with those in sympatric piscivorous fish. This data sheds light on the possibility of the utilization of periphyton as a protein-rich food by fishes.