DO IMAGE-FORMING EYES PROMOTE EVOLUTIONARY DIVERSIFICATION?

It has been suggested that image-forming eyes promote the evolutionary diversification (measured by species richness) of the groups that possess them. Several different processes could give rise to this effect, including diversifying selection in a new adaptive zone (or zones) and a reduced rate of extinction due to enhanced competitive abilities. I tested the generality of the hypothesis that imaging eyes increase net speciation by comparing extant species numbers of 12 groups that have such eyes (as categorized by Land and Fernald 1992) with those of their cladistic sister groups that lack such organs. Even assuming the published hypotheses of phylogenetic relationships that most favor increased net speciation of visual groups, these comparisons show no significant association between imaging eyes and species richness. Increased activity, as indicated by published accounts of locomotory speed, is significantly associated with the evolution of image-forming eyes. This suggests that a large "visual adaptive zone" might be characterized by relatively high activity. However, when diversity comparisons are limited to eight cases in which the evolution of imaging eyes is associated with increased activity, there is still no significant association between such eyes and species richness. The fossil record indicates that the only visual groups that have undergone major evolutionary radiations evolved imaging eyes early in the history of metazoans (before the Silurian). The radiations of these early groups may have largely filled up niches for visual animals and thus prevented the subsequent proliferation of other groups with image-forming eyes. Alternatively, it may be that image-forming eyes have no exceptional effect on diversification or that their effects are obscured by other factors in the long run.