Vrije Universiteit Brussel Genetic structure of Heliofungia actiniformis (Scleractinia: Fungiidae) populations in the Indo- Malay Archipelago: implications for live coral trade management efforts Knittweis, L.; W.S., Kraemer,; Timm, J.; Kochzius, Marc Published in: Conservation Genetics DOI: 10.1007/s10592-008-9566-5 Publication date: 2009 Document Version: Final published version Link to publication Citation for published version (APA): Knittweis, L., W.S., K., Timm, J., & Kochzius, M. (2009). Genetic structure of Heliofungia actiniformis (Scleractinia: Fungiidae) populations in the Indo-Malay Archipelago: implications for live coral trade management efforts. 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Oct. 2021 Conserv Genet (2009) 10:241–249 DOI 10.1007/s10592-008-9566-5 RESEARCH ARTICLE Genetic structure of Heliofungia actiniformis (Scleractinia: Fungiidae) populations in the Indo-Malay Archipelago: implications for live coral trade management efforts Leyla Knittweis Æ Wiebke Elsbeth Kraemer Æ Janne Timm Æ Marc Kochzius Received: 14 September 2007 / Accepted: 11 March 2008 / Published online: 8 May 2008 Ó Springer Science+Business Media B.V. 2008 Abstract The fungiid Heliofungia actiniformis is one of should be taken into consideration when setting future the most popular scleractinian coral species in the growing harvest quotas. live aquarium trade, with the majority of specimens ori- ginating in Indonesia. Details on population connectivity Keywords Mushroom coral Á Larval source Á may potentially provide important information with Genetic variation Á Larval dispersal Á Indo-Pacific regards to harvest management efforts. Genetic structure was examined, using ribosomal ITS1, 5.8S and partial ITS2 sequences on a small scale among populations in the Introduction Spermonde Archipelago, South Sulawesi (up to 65 km distance, Ust = 0.09), and on a large scale throughout the Marine ornamental species are collected around the world Indo-Malay Archipelago (up to 2,900 km distance, to supply specimens for the international aquarium trade, Ust = 0.26). Significant genetic structuring was found at which is estimated to be worth 200–330 million US$ both scales. Within the Spermonde Archipelago isolation annually. Indonesia is the world’s largest exporter of live by distance as well as local oceanographic features shaped marine ornamental corals since the 1980s (Wabnitz et al. patterns of genetic connectivity. On the large scale, the 2003) and Heliofungia actiniformis ranks in the top five of data revealed genetically distinct populations in Tomini all coral species being exploited for the live coral market Bay, New Guinea and the Thousand Islands near Jakarta, (Green and Shirley 1999). An export quota system is in and a lack of genetic differentiation among populations place, but knowledge on the population ecology of this lying close to or directly in the path of the Indonesian species is limited (Abe 1937, 1940), and altogether lacking throughflow: from the central Visayas to the Flores Sea for the Indo-Malay Archipelago. Yet if harvesting regimes (Uct = 0.32). Whilst the influence of both historical and are to be sustainable in the long term, the exploited pop- present day processes on genetic structuring of H. actini- ulations need to be self-sustaining, i.e. recruitment needs to formis populations was revealed, large scale results further exceed mortality and polyp collection. An indication of the emphasised the importance of oceanographic dynamics on extent of population connectivity provided by dispersing larval dispersal patterns in this species. Potential for larval larvae is thus important with regards to the design of input from surrounding populations, and the increased successful fishery management recommendations. vulnerability of upstream as well as isolated populations Due to the inherent difficulty in studying coral larval dispersal pathways in situ, most information to date has come from the study of the indirect effects of larval dis- L. Knittweis (&) Centre for Tropical Marine Ecology, Fahrenheitstrasse 6, persal on the genetic differentiation and gene flow among 28359 Bremen, Germany populations (e.g. Ayre and Hughes 2000; Nishikawa et al. e-mail: [email protected] 2003; Magalon 2005; Vollmer and Palumbi 2007). How- ever, a number of factors have been implicated in shaping W. E. Kraemer Á J. Timm Á M. Kochzius Biotechnology and Molecular Genetics, FB2-UFT, University present-day genetic structure of marine populations, of Bremen, Leobenerstrasse, 28359 Bremen, Germany including the roles of (1) larval dispersal capabilities (Ayre 123 242 Conserv Genet (2009) 10:241–249 and Hughes 2000; Lourie et al. 2005), (2) present day Indo-Malay Archipelago (Nelson 2000; Barber et al. 2002; physical oceanographic features (Williams and Benzie Sugama et al. 2002; Lourie et al. 2005). 1996; Rodriguez-Lanetty and Hoegh-Guldberg 2002), and Despite the central Indo-Pacific’s ecological importance (3) historical events (Nelson et al. 2000; Barber et al. as the world’s coral biodiversity hotspot (Hughes et al. 2002). 2002), and the subsistence value of local reef exploitation The dispersal potential of scleractinian coral species is to millions of people, no previous investigations into pat- influenced by the mode of sexual reproduction: in ovipa- terns of genetic variation of scleractinian coral species in rous species this involves broadcast spawning of pelagic this region are known to the authors. We used ribosomal gametes, in viviparous species the release of brooded internal transcribed spacer (ITS) sequences to (1) examine planulae. While brooded planulae generally settle within genetic variability and genetic affinities of H. actiniformis 1–2 days of release, planulae from spawning corals require populations on a small scale, i.e. throughout the Spermonde 4–6 days after gamete release before they settle (see review Archipelago (South Sulawesi) among reefs 7–65 km apart by Harrison and Wallace 1990). The only existing study on and (2) examine genetic variability and genetic affinities of the reproductive characteristics of H. actiniformis to date H. actiniformis populations on a large scale, i.e. throughout found that well developed, brooded larvae were released by the Indo-Malay Archipelago among reefs up to 2,900 km the mother polyps. The larvae showed negative phototactic apart. We discuss the implications of likely larval con- behaviour after 2 days and the great majority had attached nectivity patterns in light of potential management to the substrates provided in the laboratory after 3 days approaches to the exploitation of this species for the coral (Abe 1937). In H. actiniformis, one may thus expect aquarium trade. somewhat limited larval dispersal and potentially signifi- cant genetic structuring between neighbouring reef systems. However, it is known that coral larvae are able to Materials and methods prolong their competency periods if they do not encounter suitable substrate for settlement. Furthermore, although the Collection of samples larvae of brooding corals settle faster than those of broadcast spawners, their potential maximum competency Heliofungia actiniformis tentacle clippings were collected periods are much longer due to the high energy reserves without further moving or harming the corals from a total they can draw upon when needed (Richmond 1989; Wilson of ten sites throughout the Indo-Malay Archipelago et al. 1998). (Fig. 1). Nine locations were in Indonesia: Adi, Gilli The oceanography of the Indo-Malay Archipelago is Trawangan, Komodo, Manado, Pulau Seribu, Saboeda, dominated by the Indonesian throughflow (ITF) current, Pulau Sembilan, Spermonde Archipelago, Tilamuta and which moves up to 19 million m3 of water per second from one in the Philippines: Cebu. In the Spermonde Archipel- the Pacific to the Indian Ocean (Gordan and Fine 1996; ago samples were collected from nearshore (Barrang Gordon 2005). This current provides a potential dispersal Lompo, Samalona), midshelf (Lanyukang, Sarappokeke) corridor for marine larvae from the north-west Pacific and outershelf (Kapoposang) reefs. Wherever possible, along the Makassar Strait, all the way into the Flores and tissue samples were collected from polyps at least 50 m Banda Seas before finally being deflected to the Indian apart and of different colour morphs to reduce the likeli- Ocean. Seasonally reversing east-west currents in the Java hood of sampling genetically identical clones. Tissues were and Flores Seas moving at up to 75 cm s-1 (Wyrtki 1961) stored in 96% ethanol at 4°C. Variable numbers of samples should further facilitate present-day marine larval dispersal collected reflect species abundance levels at sampling