The dead of winter: size-dependent variation and genetic differences in seasonal mortality among Atlantic silverside (Atherinidae: Menidia menidia) from different latitudes

Abstract

Genetic differences among populations of Atlantic silverside (Menidia menidia) are hypothesized to be evolutionary responses to intense, size-selective winter mortality at high latitudes. Three experiments were conducted to test features of winter mortality. In the first experiment, we varied size and whether food was provided or withheld; temperatures were permitted to follow ambient (New York) wintertime fluctuations. Mortality and depletion of energy reserves were more rapid in the units receiving no food. Small fish died before larger fish in these units, but not in the units receiving food. Energy depletion of fish in the no-food treatment resembled that of fish in the wild. In the second experiment, we varied size and population of origin, representing high-latitude (Nova Scotia), midlatitude (New York), and low-latitude (South Carolina) populations. These fish were provided food and showed minimal depletion of energy reserves, but mortality rates were high when water temperatures were low. Mortality did not vary with size in New York and South Carolina fish, but was highest in intermediate-size fish from Nova Scotia. There was a pronounced population difference in survival rate (Nova Scotia > New York > South Carolina). In the third experiment, food was withheld and extreme low temperatures were moderated. Energy depletion was rapid and small fish died before large fish. We conclude that populations in seasonal environments are likely to be subject to size-selective winter mortality when energy reserves are depleted and that juvenile growth rates have evolved in response to this selection pressure. In addition, high-latitude populations have evolved greater tolerance to other winter stresses associated with low temperatures.