Introgression of the Gamete Recognition Molecule, Bindin, in the Sea Urchin Diadema

Hybridization is important in evolution, because it is a necessary (though not sufficient) step in the introgression of potentially adaptive variation between species. Bindin is a gamete recognition protein in echinoids and asteroids, capable of blocking cross-fertilization between species to varying degrees. Four species of the sea urchin genus Diadema are broadly sympatric in the Indo-Pacific: D. paucispinum, D. savignyi, D. clarki, and D. setosum. Data from three published studies, one of identification of hybrids through allozymes, one of the phylogeography of mitochondrial DNA, and one of the phylogeny of bindin, were combined to assess the degree of bindin introgression between these four species. I analyzed sequences of the ATPase 8 and ATPase 6 mitochondrial genes and of bindin, sampled throughout the species ranges, with an isolation-migration algorithm, IMa3. IMa3 uses a coalescent approach to produce Bayesian estimates of effective population sizes and gene flow between populations. The results showed that bindin alleles coalesce completely within the species bounds of D. clarki and of D. setosum. The sister species D. paucispinum and D. savignyi, however, were estimated as having exchanged a bindin allele at an average of every one to two-and-a-half generations since they speciated from each other. As the allozyme study detected nine hybrids between three of these species in Okinawa (most of them between D. setosum and D. savignyi) in a single sample, hybrids between these species are produced, but bindin does not introgress. Therefore, bindin must not be efficient in blocking heterospecific fertilizations. Complete, or almost complete, reproductive isolation between species of Diadema must result from low hybrid fitness.