The road toward sympatric speciation in whitefish. : The effects of divergent selection on European whitefish (Coregonus lavaretus) size and behavior, and effects on zooplankton communities.

Abstract:

For almost every organism there are large gaps in our knowledge about the processes that leads to speciation. The changes an organism undergoes before divergence has occurred have remained a mystery, as it is difficult to say whether or not a species is going to diverge and when. To investigate this unknown the European whitefish (Coregonus lavaretus) and the northern pike (Esox lucius) were studied, as they produce a repeatable and predictable pattern of speciation in sympatry. To investigate the changes in phenotypes and behaviour in whitefish that precedes divergence, two lake populations were examined, Gräsvattnet and Ringsjön. Gräsvattnet was used as a control, with a population of whitefish but an absence of pike, whereas Ringsjön has a population of whitefish that invaded from Gräsvattnet and a pike population. The presence of pike presumably exerts divergent selection on the whitefish population. Fish and zooplankton were surveyed in both lakes from 1970 to the present day, which allows us to compare how whitefish populations and their resources change in the presence and absence of pike. The results found in Ringsjön show; (1) a change in habitat use, (2) a change in diet from pelagic to benthic, (3) an increase in the relationship between individual body size and diet and (4) a decrease in average size over the course of the study. (1)The presence of pike is believed to have forced the whitefish into the pelagic which could be seen in the result, with an increase in individuals caught in the pelagic. (2) The change in diet is thought to be caused by a resource competition created by individuals being forced to use the pelagic. Although insignificant this led to an overall reduction in zooplankton abundance by almost 40% which could have intensified competition. The resource competition could then have been intensified further by the change in composition of zooplankton relative abundance. (3) The increase in relationship between individual body size and diet is thought to increase due to the resource competition between smaller and larger individuals in the pelagic. Smaller individuals are better competitors than larger individuals for pelagic resource which could have led to the larger individuals switching to a more benthic diet. (4) The decrease in average size is thought to be caused by negative selection for larger individuals. Larger individuals have switched to a more benthic diet, and although the individuals are larger they still face the risk of predation in the littoral zone as they have not outgrown the gape size of the pike. This could have led to the average size reduction that may be the first steps in speciation, and ultimately leading to the divergence of two morphs by sympatric speciation in Ringsjön. In Gräsvattnet over the course of the study there were few and small changes in whitefish size, zooplankton relative abundance in the diet and in the environment. The results in Gräsvattnet could however suggest resource competition for benthic resources. Although resource competition is thought to be an important factor in the speciation of whitefish, without predation pressure no speciation occurs. This result could suggest the importance of predation pressure in the speciation of whitefish.