Challenges for Diadromous Fishes in a Dynamic Global Environment

Abstract

<em>Abstract</em>.-We develop the view, based on life cycle differences and recently published sister group relationships, that the freshwater life cycle was the ancestral character state leading to anadromy among salmoniforms, whereas the marine life cycle was the ancestral character state leading to anadromy among osmeriforms. In contrast to most salmonid fishes, the reproductive migrations of smelts are generally characterized by brief excursions to spawn in freshwater, and larvae may spend no more than 24 h in freshwater before being transported to coastal marine or estuarine environments. We reconstructed the phylogeny of the suborder Osmeroidei to establish the phylogenetic relationships among anadromous, marine, and freshwater species of this taxon. We mapped these life cycles onto phylogenetic trees of osmeriforms and salmoniforms and applied character-reconstruction methodology based on simple parsimony and likelihood methodologies. A freshwater origin of salmonids was supported by our analyses, whereas either marine or anadromous life cycles characterized the evolution of osmeroids. The possibility that the evolution of anadromy in salmonids and osmeroids followed separate paths requires a reconsideration of some generalizations concerning anadromy. We hypothesize that anadromy in osmeroids may be first and foremost an adaptation to place embryos and the early larval stages in reproductive safe sites to maximize their survival. The evolution of exclusive freshwater species of osmeriforms has occurred via anadromy through the various processes associated with landlocking. Freshwater amphidromy in osmeroids is most likely a consequence of anadromy rather than a precursor and may be contingent upon the availability of food resources in freshwater. Finally, marine osmeroids have been derived from anadromous ancestors and are "safe-site" specialists, exploiting principally the upper intertidal zone for reproduction. We also suggest that such contrasting evolutionary pathways to anadromy may provide insight into the evolution of partial migration, observed uniquely in salmonids, and the nature and extent of population genetic structure found in the two groups of fishes.