Fish size and habitat depth relationships in headwater streams.

Surveys of 262 pools in 3 small streams in eastern Tennessee demonstrated a strong positive relationship between pool depth and the size of the largest fish within a pool (P<0.001). Similarly, the largest colonizers of newly-created deep pools were larger than the colonizers of shallow pools. We explored the role of predation risk in contributing to the "bigger fish - deeper habitat" pattern, which has been noted by others, by conducting five manipulative field experiments in two streams. Three experiments used stoneroller minnows (Campostoma anomalum); one used creek chubs (Semotilus atromaculatus); and one used striped shiners (Notropis chrysocephalus). The stoneroller experiments showed that survival of fish approximately 100 mm in total length (TL) was much lower in shallow pools (10 cm deep) than in deep (40 cm maximum) pools (19% versus 80% survival over 12 d in one experiment) and added cover markedly increased stoneroller survival in shallow pools (from 49% to 96% in an 11-d experiment). The creek chub experiment showed that, as for stonerollers, pool depth markedly influenced survival: the chubs survived an average of 4.9 d in shallow pools and >10.8 d in deep pools. In the striped shiner experiment in shallow artificial streamside troughs, no individuals 75-100 mm TL survived as long as 13 d, where-as smaller (20-25 mm) fish had 100% survival over 13 d. The results of the experiments show that predation risk from wading/diving animals (e.g., herons and raccoons) is much higher for larger fishes in shallow water than for these fishes in deeper water or for smaller fish in shallow water. We discuss the role of predation risk from two sources (piscivorous fish, which are more effective in deeper habitats, and diving/wading predators, which are more effective in shallow habitats) in contributing to the bigger fish - deeper habitat pattern in streams.