We report here, the effects of extended competency on larval survival, metamorphosis, and postlarval juvenile growth of four closely related species of tropical sea urchins, Echinometra sp. A (Ea), E. mathaei (Em), Echinometra sp. C (Ec), and E. oblonga (Eo). Planktotrophic larvae of all four species fed on cultured phytoplankton (Chaetoceros gracilis) attained metamorphic competence within 22-24 days after fertilization. Competent larvae were forced to delay metamorphosis for up to 5 months by preventing them from settling in culture bottles with continuous stirring on a set of 10 rpm rotating rollers and larval survival per monthly intervals was recorded. Larval survival was highest at 24 days, when competence was attained (0 delayed period), and there were no significant differences among the four species. Larvae that had experienced a prolonged delay had reduced survival rate, metamorphosis success, and juvenile survival, but among older larvae, Em had the highest success followed by Ea, Eo, and Ec. Juveniles from larvae of all four species that metamorphosed soon after becoming competent tended to have higher growth rates (test diameter and length of spines) than juveniles from larvae that metamorphosed after a prolonged period of competence with progressively slower growth the longer the prolonged period. Despite the adverse effects of delaying metamorphosis on growth parameters, competent larvae of all four species were able to survive up to 5 months and after metamorphosis grew into 1-month-old juveniles in lab condition. Overall, delayed larvae of Em showed significantly higher larval survival, metamorphosis, and juvenile survival than Ea and Eo, while Ec showed the lowest values in these performances. Em has the most widespread distribution of these species ranging from Africa to Hawaii, while Ec probably has the most restricted distribution. Consequently, differences in distribution may be related to differences in the ability to delay metamorphosis.
Two reef margin species of tropical sea urchins, Echinometra sp. C (Ec) and Echinometra oblonga (Eo), occur sympatrically on Okinawa intertidal reefs in southern Japan. Hybridization between these species was examined through a series of cross-fertilization experiments. At limited sperm concentrations, where conspecific crosses reached near 100% fertilization, both heterospecific crosses showed high fertilization rates (81%-85%). The compatibility of the gametes demonstrated that if gamete recognition molecules are involved in fertilization of these species, they are not strongly species-specific. We found that conspecific crosses reached peak fertilization levels much faster than did heterospecific crosses, indicating the presence of a prezygotic barrier to hybridization in the gametes. Larval survival, metamorphosis, and juvenile and adult survival of hybrid groups were nearly identical to those of their parent species. Hybrids from crosses in both directions developed normally through larval stages to sexually mature adults, indicating that neither gametic incompatibility nor hybrid inviability appeared to maintain reproductive isolation between these species. In adults, Ec×Ec crosses gave the highest live weight, followed by Eo (ova)×Ec (sperm), Ec (ova)×Eo (sperm), and Eo×Eo. Other growth performance measures (viz., test size, Aristotle's lantern length, and gonad index) of hybrid groups and their parental siblings showed the same trends. The phenotypic color patterns of the hybrids were closer to the maternal coloration, whereas spine length, tube-foot and gonad spicule characteristics, pedicellaria valve length, and gamete sizes showed intermediate features. Adult F(1) hybrids were completely fertile and displayed high fertilization success in F(1) backcrosses, eliminating the likelihood that hybrid sterility is a postzygotic mechanism of reproductive isolation. Conversely, intensive surveys failed to find hybrid individuals in the field, suggesting the lack or rarity of natural hybridization. This strongly suggests that reproductive isolation is achieved by prezygotic isolating mechanism(s). Of these mechanisms, habitat segregation, gamete competition, differences in spawning times, gametic incompatibility or other genetic and non-genetic factors appear to be important in maintaining the integrity of these species.