Evidence for general size-by-habitat rules in actinopterygian fishes across nine scales of observation

Identifying environmental predictors of phenotype is fundamentally important to many ecological questions, from revealing broadscale ecological processes to predicting extinction risk. However, establishing robust environment-phenotype relationships is challenging, as powerful case studies require diverse clades which repeatedly undergo environmental transitions at multiple taxonomic scales. Actinopterygian fishes, with 32,000+ species, fulfil these criteria for the fundamental habitat divisions in water. With four datasets of body size (ranging 10,905-27,226 species), I reveal highly consistent size-by-habitat-use patterns across nine scales of observation. Taxa in marine, marine-brackish, euryhaline and freshwater-brackish habitats possess larger mean sizes than freshwater relatives, and the largest mean sizes consistently emerge within marine-brackish and euryhaline taxa. These findings align with the predictions of seven mechanisms thought to drive larger size by promoting additional trophic levels. However, mismatches between size and trophic-level patterns highlight a role for additional mechanisms, and support for viable candidates is examined in 3439 comparisons.