Symbiosis as an evolutionary innovation in the radiation of Paleocene planktic foraminifera

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Abstract
Symbioses are often regarded as an important means for the creation of evolutionary novelty as well as a trigger for the abrupt appearance of higher taxa. The fossil record of foraminifer-algal symbiosis suggests that the appearance of this ecological association contributed to the radiation of Paleogene planktic foraminifera. Isotopic evidence shows that photosymbiosis evolved in synchrony with a major diversification of trochospiral planktic foraminifera about 3.5 m.y. after the end-Cretaceous extinction. In modern planktic foraminifera, photosymbiotic species tend to have more cosmopolitan distributions than asymbiotic foraminifera and a greater ability to withstand periods of nutrient stress. The simultaneous taxonomic radiation and acquisition of photosymbiosis are evidence that the ecological strategy permitted Paleocene foraminifera to expand their niche in pelagic environments by diversifying into low-nutrient surface waters.A comparison of the species longevities of Neogene and Paleogene symbiotic clades suggests that photosymbiosis does not regulate the characteristic rate of taxonomic turnover in clades after they appear. Species longevities are much shorter in Paleocene and Eocene photosymbiotic morphospecies than they are among photosymbiotic Neogene clades; apparently photosymbiosis does not exert a significant control over long-term evolutionary rates. In addition, the absence of a characteristic morphology associated with photosymbiosis in Cenozoic planktic foraminifera suggests that morphology, as with rate of evolutionary turnover, is linked to symbiosis only because of common inheritance instead of a functional relationship. Although the coincidence between the acquisition of photosymbiosis and generic diversification does suggest a linkage between this ecology and the appearance of foraminiferal higher taxa, there is little indication at the present that symbioses control long-term morphological or ecological patterns within these groups after their appearance. Photosymbiosis, and other evolutionary innovations, may be more a catalyst for the differentiation of major groups than a predictable governor on evolutionary rates.