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Exploring the Role of Micronesian Islands in the Maintenance of Coral Genetic Diversity in the Pacific Ocean

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Exploring the Role of Micronesian Islands in the Maintenance of Coral Genetic Diversity in the Pacific Ocean Molecular Ecology (2015) 24, 70–82 doi: 10.1111/mec.13005 Exploring the role of Micronesian islands in the maintenance of coral genetic diversity in the Pacific Ocean S. W. DAVIES,* E. A. TREML,† C. D. KENKEL* and M. V. MATZ* *Department of Integrative Biology, The University of Texas at Austin, 1 University Station C0990, Austin, TX 78712, USA, †Department of Zoology, University of Melbourne, Melbourne, Vic. 3010, Australia Abstract Understanding how genetic diversity is maintained across patchy marine environments remains a fundamental problem in marine biology. The Coral Triangle, located in the Indo-West Pacific, is the centre of marine biodiversity and has been proposed as an important source of genetic diversity for remote Pacific reefs. Several studies highlight Micronesia, a scattering of hundreds of small islands situated within the North Equa- torial Counter Current, as a potentially important migration corridor. To test this hypothesis, we characterized the population genetic structure of two ecologically important congeneric species of reef-building corals across greater Micronesia, from Palau to the Marshall Islands. Genetic divergences between islands followed an isola- tion-by-distance pattern, with Acropora hyacinthus exhibiting greater genetic diver- gences than A. digitifera, suggesting different migration capabilities or different effective population sizes for these closely related species. We inferred dispersal dis- tance using a biophysical larval transport model, which explained an additional 15– 21% of the observed genetic variation compared to between-island geographical dis- tance alone. For both species, genetic divergence accumulates and genetic diversity diminishes with distance from the Coral Triangle, supporting the hypothesis that Micronesian islands act as important stepping stones connecting the central Pacific with the species-rich Coral Triangle. However, for A. hyacinthus, the species with lower genetic connectivity, immigration from the subequatorial Pacific begins to play a larger role in shaping diversity than input from the Coral Triangle. This work high- lights the enormous dispersal potential of broadcast-spawning corals and identifies the biological and physical drivers that influence coral genetic diversity on a regional scale. Keywords: Acropora, biophysical model, dispersal, genetic diversity, isolation by distance, stepping stone Received 8 August 2014; revision received 4 November 2014; accepted 7 November 2014 quences for the persistence, speciation and extinction of Introduction numerous marine taxa. Understanding the limitations Waters of the Indo-West Pacific (also termed ‘The Coral to gene flow to and from the Coral Triangle will Triangle’) support the greatest tropical marine biodiver- improve predictions of how marine biodiversity pat- sity on the planet (Briggs 1987; Vernon 1995; Veron terns might be affected by global climate change (Bur- et al. 2009). The processes responsible for generating rows et al. 2011), which is invaluable both for and redistributing this diversity have significant conse- prioritization of management efforts and basic under- standing of evolution in the ocean. Correspondence: Sarah W. Davies, Fax:512 471 3878; Several models of seascape connectivity between the E-mail: [email protected] Coral Triangle and its surrounding oceans have been © 2014 John Wiley & Sons Ltd CORAL GENETIC CONNECTIVITY ACROSS MICRONESIA 71 developed (Treml et al. 2008; Kool et al. 2011; Wood spawning characteristics. Importantly, the group of et al. 2014). Most recently, Wood et al. (2014) inferred organisms that serve as the foundation for these ecosys- little direct migration from the Coral Triangle to the tems, reef-building corals (phylum Cnidaria, class central Pacific, but instead proposed that the Coral Tri- Anthozoa, order Scleractinia), remain understudied in angle supplies larvae to Micronesia, which in turn the Pacific (Keyse et al. 2014). serves as a source for the central Pacific via the North Coral dispersal is potentially extensive because most Equatorial Counter Current (NECC, Fig. 1). Micronesia major reef-building species reproduce by releasing is therefore hypothesized to serve as a series of step- gametes into the water column, resulting in pelagic ping stones connecting the Coral Triangle to the central larvae that disperse broadly with ocean currents (Baird Pacific. If species from the Coral Triangle are incapable et al. 2009). Many larvae can survive for months in the of dispersing the expanse of Micronesia within one gen- absence of settlement cues (Graham et al. 2008, 2013), eration, differentiation in gene frequencies should build and different species appear to be flexible in the length up across Micronesia as distances between populations of their pelagic larval duration (PLD) (Connolly & increase (i.e. an ‘isolation-by-distance (IBD)’ pattern Baird 2010). Coral population genetic studies have should be observed) (Wright 1943; Kimura & Weiss revealed gene flow on scales ranging from tens to hun- 1964; Slatkin 1993). This prediction can be explicitly dreds of kilometres (Ayre & Hughes 2000, 2004; tested through population genetic methods. Underwood et al. 2009; Torda et al. 2013), with evi- Few studies have explored genetic connectivity dence for long-distance dispersal (Baums et al. 2005; within Micronesia and between Micronesia and the rest Severance & Karl 2006; van Oppen et al. 2011), result- of the Pacific, and thus far dispersal patterns remain ing in high genetic connectivity and large geographical unresolved. A study of the yellow tang fish (Zebrasoma ranges for many coral species. However, recent work flavescens) found support for westward migration from suggests that dispersal distances for many taxa are less Hawaii to the Central Pacific (Pohnpei) (Eble et al. than previously assumed. There is high potential for 2011), contrasting with the eastward migration patterns local retention, where individuals remain in the same suggested in studies of other marine species (Priest population from which they originated (Figueiredo et al. 2012; Timmers et al. 2012). This lack of consistency et al. 2013), suggesting that wide-ranging species may in dispersal patterns among taxa might be due to varia- rely on stepping stones for long-distance gene flow. tions in species life history, dispersal capabilities or Given the extensive ranges for most coral species, it is Fig. 1 Geographic locations of the Micro- nesian islands where A. hyacinthus and A. digitifera corals were collected. Top: map of the Micronesian Pacific with an inset of the Pacific Ocean for reference. Islands where samples were collected are designated with grey boxes. The subset of islands included in the biophysical model are marked with an (*). Coloured blocks are estimates of dominant current patterns (Bonjean & Lagerloef 2002). Yel- low designates the North Equatorial Cur- rent (NEC). Pink designates the North Equatorial Counter Current (NECC). Blue represents the South Equatorial Current (SEC). Arrows in each quadrant repre- sent current direction. Bottom: enlarged regional maps for each island with sam- pling sites shown in boxes. Detailed information on each sampling site is located in Table 1. © 2014 John Wiley & Sons Ltd 72 S. W. DAVIES ET AL. reasonable to predict that Micronesian islands serve as Materials and methods effective stepping stones, even though these islands are small, remote and separated by large expanses of open Detailed materials and methods can be found in ocean. Appendix S1 (Supporting information). Here, we implemented a seascape genetics study of two Pacific coral species of the genus Acropora, the Sampling locations and methodology most species-rich and ecologically important genus with some of the largest geographical ranges [i.e. From 2009 to 2011, snorkelling or scuba (3–7 m depth) A. digitifera range >100 000 km2 (Veron 2000)]. We sam- was used to sample 23 reef sites on ten islands through- pled A. digitifera and A. hyacinthus over the entire out Micronesia (Fig. 1, Table 1). Fifty colonies (>2m range of greater Micronesia, in addition to one subequ- apart) of each coral species (Acropora hyacinthus and atorial Pacific location to test (i) what are the dispersal A. digitifera) were sampled per reef. Both species were limits for a coral with a long PLD and a large geo- collected at the same GPS coordinates and depth was graphical range; (ii) whether the distribution of regio- maintained constant within a sampling site whenever nal genetic diversity is consistent with a stepping stone possible. Colonies were photographed for confirmation model; (iii) to which extent biophysical models of lar- of species identification, and one small (~2 cm), ran- val dispersal can explain observed genetic differentia- domly chosen branch tip was collected, preserved in tion patterns; (iv) whether two closely related species 96% ethanol and stored at À20 °C. A total of 2095 speci- sharing the same reproductive strategy exhibit similar mens were collected. An additional site south of the genetic structures; and (v) whether Micronesia serves equator (Phoenix Islands N = 19 collected in 2008) was as a dispersal corridor between the Coral Triangle and included post hoc to test specific dispersal hypotheses the central Pacific. for A. hyacinthus. Table 1 Reef Site Collections. GPS coordinates, main island group, and number of A. digitifera and A. hyacinthus genotyped. Site let- ter corresponds to Fig. 1 island insets Site Island GPS A. digitifera A. hyacinthus a. West Channel Reef Palau 7°31055.7 N, 134°29042.8 E 39 44 b. Lighthouse Reef Palau 7°16062.4 N, 134°27061.9 E 49 50 c. Ngulu Ngulu Atoll 8°18012.0 N, 137°29018.7 E 02 46 d. South Tip Reef Yap 9°26005.4 N, 138°02010.4 E 45 48 e. West Outer Reef Yap 9°33047.3 N, 138°05071.5 E 46 50 f. Goofnuw Channel Yap 9°34026.4 N, 138°12019.2 E 49 37 g. Pago Bay Guam 13°25066.6 N, 144°47094.3 E 45 0* h. Tanguisson Guam 13°32061.1 N, 144°48052.6 E 50 0* i. West Polle Chuuk 7°19069.7 N, 151°33021.1E 451 39 j. Aroch Patch Reef Chuuk 7°14042.0 N, 151°53095.4 E 02 49 k.
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