Intraspecific diversity and ecological zonation in coral-algal symbiosis.

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Abstract
All reef-building corals are obligately associated with photosynthetic microalgal endosymbionts called zooxanthellae. Zooxanthella taxonomy has emphasized differences between species of hosts, but the possibility of ecologically significant zooxanthella diversity within hosts has been the subject of speculation for decades. Analysis of two dominant Caribbean corals showed that each associates with three taxa of zooxanthellae that exhibit zonation with depth--the primary environmental gradient for light-dependent marine organisms. Some colonies apparently host two taxa of symbionts in proportions that can vary across the colony. This common occurrence of polymorphic, habitat-specific symbioses challenges conventional understanding of the units of biodiversity but also illuminates many distinctive aspects of marine animal-algal associations. Habitat specificity provides ecological explanations for the previously documented poor concordance between host and symbiont phylogenies and the otherwise surprising lack of direct, maternal transmission of symbionts in many species of hosts. Polymorphic symbioses may underlie the conspicuous and enigmatic variability characteristic of responses to environmental stress (e.g., coral "bleaching") and contribute importantly to the phenomenon of photoadaptation.