The evolution of fish ectoparasite communities--the role of the ice ages.

The monogenean ectoparasite genera Dactylogyrus and Gyrodactylus are hyperdiverse, and yet monogenean communities infecting freshwater fish in the northern Holarctic are variable and often impoverished. This is a result of extinctions during periods of glaciation when the host fishes experienced range contraction. Fish species recolonised northern Europe by one of three routes; stenohaline fishes recolonised from the Black Sea basin via Danube-Rhine or Dnieper-Dniester-Vistula connections. These fish retain complex rich monogenean faunas with numerous species of Dactylogyrus and Paradiplozoon. Anadromous fishes, recolonising via marine migrations along the coast, and cold-water fishes surviving in refugia close to the ice sheets, have impoverished monogenean faunas, characterised by gyrodactylid and ancyrocephalid species. However, there may be great complexity, due to the evolution of new host-parasite interactions within specific watersheds before, during and after the Last Glacial Maximum (LGM). This is demonstrated using examples from the sticklebacks (Gasterosteus aculeatus) and the bullheads (Cottus gobio). Finally, the evolution of Gyrodactylus salaris from a clade of G. thymalli is described, and the role of glaciation in stimulating the evolution of numerous clades of the latter is discussed. This latter example represents a unique opportunity to study speciation by host shift in real time.