Persistence of Corals in Marginal Habitats: the Role of the Environment, and Symbiont Diversity and Ecophysiology
Many corals live in marginal habitats, close to their survival thresholds of water temperature, light penetration and aragonite saturation. Living under these highly variable and extreme conditions is likely facilitated by specific physiological adaptations and/or the presence of unique species of coral and their symbionts but data on these factors are limited. The specific objectives of the study were to: (1) examine the diversity and distribution patterns of corals in marginal environments, (2) investigate the diversity, distribution patterns and host specificity of symbionts in corals in marginal environments, (3) assess the influence of environmental variables on host and symbiont distribution in marginal environments, in comparison to 'optimal' environments, and (4) examine the physiological responses to changing environmental conditions and stress of corals and their symbionts in marginal environments. Surveys of coral community patterns were conducted at the Kermadec Islands (KI), New Zealand, and Palmyra Atoll, USA, with local scale environmental parameters (i.e. wave exposure and sedimentation) found to control the diversity and distribution of the coral communities. Symbiodinium types were identified to subcladal level in a range of coral species at each of the survey sites, using ITS2-DGGE. A high diversity of C type symbionts (19 types in 13 host genera), and reduced host specificity was observed at the high latitude site of Lord Howe Island (LHI), Australia, with similarly high diversity at the KI (10 types in 9 genera). Thirteen novel clade C types were identified in corals at LHI, with two of these types also present in hosts at the KI. The reduced host specificity of symbionts at LHI, compared to tropical sites, implies that the evolution of novel holobionts may be an important mechanism whereby corals can cope with variable and stressful conditions. Further, physiological assessment of the novel LHI symbionts led to the suggestion that Symbiodinium at LHI may be specialised for cooler and more variable temperatures, so contributing to the success of corals at this marginal location. In contrast, a low diversity of generalist symbionts (C and D types) were uncovered at the equatorial site of Palmyra Atoll (10 types in 13 genera), attributed to the stressful environmental regime resulting in a reduced population of stresstolerant symbionts. The variation in environmental parameters, particularly sedimentation, around Palmyra Atoll has led to diversification of coral communities, however this environmental variation has not affected the symbiont communities. While it has been suggested that marginal coral communities might be better adapted for survival in an environment modified by global climate change, the local scale environmental factors are also important drivers of both coral and symbiont distributions, and should be considered when making predictions for the future. Further, assessment of the physiological tolerance ranges of both the multiple, novel symbionts at high latitudes, and the few, potentially stress-tolerant symbionts at Palmyra should be conducted, to help determine whether they have the ability to adjust to new environmental conditions.