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Gene Flow in Coral Reef Organisms of the Tropical Eastern Pacific 16 Gene Flow in Coral Reef Organisms of the Tropical Eastern Pacific 16 H.A. Lessios and Iliana B. Baums Abstract Gene flow can provide cohesion between conspecific populations. In order to obtain an indirect measure of gene flow between coral reef species in the eastern tropical Pacific (ETP) and between these populations and those of the rest of the Pacific we compiled available data from sequences of DNA and microsatellites for corals, gastropods, echinoderms and fishes, and calculated FST statistics. The ETP consists of a narrow strip of continental shelf along the coast of the Americas and a deeper water gap between the coast and the outer eastern Pacific Islands; a large expanse of deep ocean separates the ETP and the closest islands in the central Pacific. We have, therefore, compared populations in four major directions: (1) between the eastern and the central Pacific, (2) between the coast and the outer islands, (3) among the outer islands, and (4) along the coast and nearshore islands. The available data are biased in favor of showing high levels of gene flow because they contain an excess of transpacific species, which are a minority among ETP biota. Despite this bias, shallow water populations of the ETP are isolated from the rest of the world’s oceans. Occasional breaching of the expanse of water between the ETP and the Central Pacificby some species is also possible. Gene flow between the outer eastern Pacific islands and the mainland coast is variable, depending on the species examined. Gene flow among populations at the outer eastern Pacific islands is high except for those at Easter Island (Rapa Nui), in which all but one sampled species show large and significant values of FST in comparisons with populations from all other islands. Gene flow rates among populations along the ETP coast are high. There is no evident genetic break resulting from the Central American Gap (southern Mexico to the Gulf of Fonseca, Honduras) in any of the sampled species. A trend of isolation by distance along the coast is evident in corals and fishes. Keywords Genetic connections Á Mitochondrial DNA Á Microsatellites Á Genetic structure Á FST statistics 16.1 Introduction Gene flow is the exchange of genes between conspecific H.A. Lessios (&) populations. It is an important biological process in every Smithsonian Tropical Research Institute, 0843-03092 Balboa, species because gene transfer provides cohesion between Panama populations, preventing their independent evolution. The e-mail: [email protected] movement of genes over generations assures that new I.B. Baums advantageous mutations that arise in one location can The Pennsylvania State University, 208 Mueller Laboratory, eventually spread throughout the species’ range. When gene University Park, PA 16802, USA fl “ e-mail: [email protected] ow becomes restricted populations become genetically © Springer Science+Business Media Dordrecht 2017 477 P.W. Glynn et al. (eds.), Coral Reefs of the Eastern Tropical Pacific, Coral Reefs of the World 8, DOI 10.1007/978-94-017-7499-4_16 478 H.A. Lessios and I.B. Baums structured”, diverging from each other and (if the restrictions 16.2 Oceanographic Features become severe and last a long time) possibly turning into of the Tropical Eastern Pacific separate species. The sharing of genes between populations Relevant to Gene Flow can also dilute adaptation to local environmental conditions, thus averaging the response to natural selection over the The ETP (Fig. 16.1) consists of a narrow strip of continental connected populations. It may thus limit the geographic shelf along the coast of the American continent with some extent over which a species can spread, as peripheral pop- islands close to the shore, a deeper water gap between the ulations are prevented from adapting to conditions at the coast and the outer eastern Pacific Islands (Easter Island, the edge of a species range and then proceed to colonize new, Galápagos Archipelago, Malpelo Island, Isla del Coco, previously unsuitable, areas (Haldane 1956; Case and Taper Clipperton Atoll and the Revillagigedo Archipelago). 2000; Sexton et al. 2009; Dawson et al. 2010). A large expanse of deep ocean separates the ETP and the Genetic exchange is effected by the dispersal of propag- closest islands in the central Pacific (the Line Islands, the ules from one population to another, but, as propagules are Marquesas, and Hawaii). Thus, shallow water populations in difficult to follow, most estimates of gene flow do not the ETP can exchange genes in four major directions: involve direct observations of such transfers; they rely (1) between the eastern and the central Pacific, (2) between instead on assessing divergence between populations. This is the mainland coast and the outer islands, (3) among the outer particularly true for marine organisms in which planktonic islands, and (4) along the coast and nearshore islands. larvae maintain genetic connections over very long distances Potential barriers to gene flow in each direction consist of and are virtually impossible to track in the water column. habitat unsuitable for the establishment of adult populations Species with this kind of larvae are expected to show high and of currents that channel the movement of larvae along a rates of genetic exchange between populations and thus high particular vector, or that alter environmental conditions as to genetic homogeneity, although this expectation is not always exceed the tolerances of propagules in a particular area (see met. Several studies have indicated that in a number of Chaps. 3 and 4, Fiedler and Lavín, and Wang et al., marine species self-recruitment to the natal population is respectively). Expanses of deep water that are difficult to high, and the dispersal potential expected of their larvae fails cross in a single larval life can be expected to cause the most to be realized (Jones et al. 1999; Swearer et al. 1999, 2002; marked restrictions to the exchange of genes. If impassable, Hellberg et al. 2002; Taylor and Hellberg 2003). Such these barriers can prevent the spread of species, thus estab- studies have led to doubts that the dogma of near-panmixia lishing different biogeographic provinces as determined by in marine organisms is ever realized (Hellberg 2009). Even patterns of species presence and absence. when local recruitment does occur, however (as it Physical barriers that impede gene exchange between undoubtedly does in most populations), and even if the populations in the ETP and those in the rest of the world fraction of the larvae that recruit successfully at a distant define it as a separate oceanic region. Towards the east, an locality is small, it may be sufficient to homogenize genetic uninterrupted land bridge has existed for the last 2–3 million constitution of populations over large distances. years (Coates and Obando 1996). [It has been claimed The position of coral reef organisms of the eastern tropical recently that a nearly complete barrier to water exchange Pacific (ETP) in the spectrum between panmixia and high with the Caribbean has existed since the Eocene or early genetic structure is the subject of this chapter. We begin by Miocene (Montes et al. 2012) but this claim is incompatible considering geographical and oceanographic features that with paleoceanographic (Keigwin 1982; Collins 1996; Haug would be expected to affect rates of contemporary and his- and Tiedemann 1998;O’Dea et al. 2007), paleontological torical gene flow, then summarize existing data from the (Webb 1976; Coates and Obando 1996), and genetic (Les- literature about genetic connectivity. Not all of the data were sios 2008) data.] Towards the west lies the widest marine gathered to assess population structure, and the organisms for biogeographical barrier on the planet, 4000–7000 km of which genetic data exist are far from uniformly spread over deep water without any stepping stone habitats on which taxonomic groups. Nevertheless, a compilation of such data adults of shallow water marine species can exist (Ekman can begin to address the question of whether general patterns 1953; Briggs 1974). This ocean configuration, known as the exist. In “coral reef organisms” we include all species that are “Eastern Pacific Barrier” (EPB) has been in place for most of represented by populations resident in coral reefs, even if the Cenozoic (Grigg and Hey 1992). It is so difficult to cross, they are not exclusively found in this particular habitat. As that most shallow water benthic genera are represented by most of these species (even corals) are capable of living different species on its two sides. Had it not been for the outside coral reefs (Robertson 1998; Guzmán et al. 2004), small number of “transpacific species”, species that span the this means that we have attempted to include studies on all EPB (Emerson 1978, 1982; Vermeij 1978; Rosenblatt and organisms that live on hard substrata in the photic zone, Waples 1986; Lessios et al. 1996; Robertson et al. 2004; except for those limited to the upper intertidal shore line. Lessios and Robertson 2006), it would not have figured in a 16 Gene Flow in Coral Reef Organisms of the Tropical Eastern Pacific 479 Fig. 16.1 Ten year mean (1993– 2003) ocean surface currents in the Central-East Pacific(a) and the Eastern Tropical Pacific(b). Given are surface current vectors with 1 degree (a) or 1/3 degree (b) resolution. Basemap generated with ETOPO 2 (USGS). CC = California Current, NEC = North Equatorial Current, NECC = North Equatorial Counter Current, SEC = South Equatorial Current, CRCC = Costa Rica Coastal Current discussion of conspecific gene flow. The barrier is traversed species from the species-rich central Pacific into the ETP by the westerly North and South Equatorial Currents and by (Dana 1975; Richmond 1990; Robertson et al. 2004) and the easterly North Equatorial Counter Current.
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