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Research Article Reef Fish Dispersal in the Hawaiian Archipelago: Comparative Phylogeography of Three Endemic Damselfishes

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Research Article Reef Fish Dispersal in the Hawaiian Archipelago: Comparative Phylogeography of Three Endemic Damselfishes Hindawi Publishing Corporation Journal of Marine Biology Volume 2016, Article ID 3251814, 17 pages http://dx.doi.org/10.1155/2016/3251814 Research Article Reef Fish Dispersal in the Hawaiian Archipelago: Comparative Phylogeography of Three Endemic Damselfishes Kimberly A. Tenggardjaja,1 Brian W. Bowen,2 and Giacomo Bernardi1 1 Department of Ecology and Evolutionary Biology, University of California Santa Cruz, 100 Shafer Road, Santa Cruz, CA 95060, USA 2Hawaii Institute of Marine Biology, University of Hawaii, P.O. Box 1346, Kaneohe, HI 96744, USA Correspondence should be addressed to Kimberly A. Tenggardjaja; [email protected] Received 2November 2015; Accepted 8March 2016 Academic Editor: Yehuda Benayahu Copyright © 2016 Kimberly A. Tenggardjaja et al. Tis is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Endemic marine species at remote oceanic islands provide opportunities to investigate the proposed correlation between range size and dispersal ability. Because these species have restricted geographic ranges, it is assumed that they have limited dispersal ability, which consequently would be refected in high population genetic structure. To assess this relationship at a small scale and to determine if it may be related to specifc reef fsh families, here we employ a phylogeographic survey of three endemic Hawaiian damselfshes: Abudefduf abdominalis, Chromis ovalis,andChromis verater. Data from mitochondrial markers cytochrome b and control region revealed low but signifcant genetic structure in all three species. Combining these results with data from a previous study on Dascyllus albisella and Stegastes marginatus,allfveendemicdamselfshspeciessurveyedtodateshowevidence of genetic structure, in contrast with other widespread reef fsh species that lack structure within the Hawaiian Archipelago. Tough individual patterns of connectivity varied, these fve species showed a trend of limited connectivity between the atolls and low-lying Northwestern Hawaiian Islands versus the montane Main Hawaiian Islands, indicating that, at least for damselfshes, the protected reefs of the uninhabited northwest will not replenish depleted reefs in the populated Main Hawaiian Islands. 1. Introduction usually the products of long periods of isolated local recruit- ment and reproduction, they serve as model study organisms Due to an apparent lack of barriers in the ocean and the for understanding dispersal [5]. Te general assumption potential for larvae to disperse long distances via ocean has been that the constrained geographic range sizes of currents, the previously long-held paradigm has been that endemic species refect limited dispersal abilities [9,12,13], there is abundant connectivity and consequently little genetic yet retention-favorable traits are not common characteristics diferentiation between populations of marine organisms of ocean island endemics [5,14]. For instance, pelagic larval [1–3]. However, studies demonstrating self-recruitment and duration (PLD) is a life history trait that provides an intuitive local larval retention indicate that not all marine organisms gauge of dispersal, by the logic that more time spent in the are exhibiting broad-scale larval dispersal [4–7]. In these plankton results in greater dispersal and connectivity [15–17]. circumstances, research has shifed toward understanding However, endemic reef fshes do not show a trend toward the factors mediating connectivity in marine systems and shorter PLDs relative to widespread congeners, and some whether there are general patterns related to phylogenetic studies have shown the opposite [9,14,18]. groups, pelagic larval duration, ecology, or behavior [8–11]. While no diagnostic life history traits related to Nevertheless, generalizations have proven elusive. endemism have been identifed, there is support for a positive Isolated oceanic islands provide an excellent opportunity correlation between dispersal ability and range size [19,20]. for investigating dispersal in marine organisms. Rates of Eble et al. [21] sought to evaluate this relationship through a endemism are markedly high, and since endemic species are phylogeographic comparison in the Hawaiian Archipelago 2 Journal of Marine Biology of three surgeonfshes (family Acanthuridae) with diferent structure, we would predict genetic diferentiation across the geographic ranges. Te Hawaiian endemic was predicted to ranges of A. abdominalis, C. ovalis,andC. verater as well, exhibit less genetic connectivity (more genetic structure) providing more support for a correlation between range size than widespread members of the family. Results supported and dispersal ability. Additionally, this fnding may indicate this hypothesis, with the endemic species demonstrating that genetic diferentiation is typical of endemic Hawaiian more, albeit weak, genetic structure than the two species with damselfshes. broader geographic distributions. In the Galapagos Islands, Results from our study also contribute to the conser- Bernardi et al. [22] surveyed reef fsh species with varying vation of the Hawaiian Archipelago. Te NWHI host the range sizes, and again the endemic species demonstrated less Papahanaumoku¯ akea¯ Marine National Monument, one of genetic connectivity than species with broader distributions. the largest marine protected areas in the world and the Likewise, in a meta-analysis of tropical reef fshes, the largest in the US. Te degree of connectivity between the relationship between range size and dispersal potential, as NWHI and the MHI is of particular interest to the man- inferred from PLD, was found to vary between oceans, with a agement of marine resources in the archipelago. Te vast signifcant correlation demonstrated in the Indo-Pacifc [20]. and uninhabited marine protected area (NWHI), adjacent Tis relationship strengthened at higher taxonomic levels to a large community that depends on the sea for nutrition and was signifcant in the damselfshes (Pomacentridae), (MHI), is postulated to have a spillover efect [35,36]. Our wrasses (Labridae), and butterfyfshes (Chaetodontidae), fne-scale sampling throughout the Hawaiian Islands can indicating that phylogenetic afliation is a component of this illustrate whether the NWHI have the potential to subsidize relationship. the overexploited reefs of the MHI. Here we assess genetic connectivity across the Hawaiian Archipelago, which is one of the most isolated archipelagoes 2. Materials and Methods in the world and has 25% endemism for shore fshes [23, 24].Te archipelago comprises eight Main Hawaiian Islands 2.1. Tissue Collection. Collections of 345 A. abdominalis,412 (MHI), which are “high islands” of volcanic basaltic com- C. ovalis,and425C. verater specimens (fn clips) were made position, and ten Northwestern Hawaiian Islands (NWHI), at 13–15 locations across the Hawaiian Archipelago from 2009 which are mostly “low islands” with coral reefs and sand to 2012 (Figure 1). Additional C. verater specimens were banks overgrowing subsided basaltic foundations [25]. In this collected at Johnston Atoll ( ). Collections were made study, we focused on three endemic Hawaiian damselfshes: with pole spears or hand nets while snorkeling or SCUBA Abudefduf abdominalis, Chromis ovalis,andChromis verater. diving. #=47 Tese three species have ranges that span the entire Hawaiian Archipelago, and C. verater is also found at Johnston Atoll, about 860km south of Hawaii. Johnston Atoll is part of the 2.2. DNA Extraction, Marker Amplifcation, and Sequencing. Hawaiian marine biogeographic province because its marine Tissue specimens were preserved in salt-saturated water with fauna is predominantly Hawaiian [26]. Hence, species that 20% DMSO [37]. All of the protocols for DNA extraction, only occur in the Hawaiian Archipelago and Johnston Atoll marker amplifcation, and sequencing are identical to those are still regarded as Hawaiian endemics. used in Tenggardjaja et al. [33]. Cytb and mitochondrial Our study is preceded by a survey of two endemic Hawai- CR sequences of C. verater generated for Tenggardjaja et ian damselfshes: Stegastes marginatus and Dascyllus albisella al. [33] were used in this study. Additionally, since the lab [27]. Ramon et al. [27] analyzed the mitochondrial control work for the current study was conducted concurrently with region (CR) and found genetic structure in both species, the study on A. abdominalis by Coleman et al. [38], cytb in contrast to the majority of reef fshes surveyed across sequences of A. abdominalis were shared between the authors. Hawaii,whichshownostructurewithinthearchipelagousing Of these sequences, thirteen were identifed as hybrids by the mitochondrial marker cytochrome b (cytb) [28–31] (but Coleman et al. [38] and were included in the current study see [32]). Furthermore, one of our study species, C. verater, afer determining that they did not bias mtDNA analyses. was the subject of a separate study on connectivity between Sequences were aligned using the Geneious aligner and shallow and mesophotic ( 30 m) reef habitats [33]. No ver- edited using GENEIOUS R6 (Biomatters, LTD, Auckland, tical (depth-related) structure was identifed in this species, NZ). Alignments of cytb were unambiguous, while CR contained multiple indels of 1-2bp. Unique haplotypes for but the Hawaiian Archipelago> was signifcantly diferentiated each marker were identifed in ARLEQUIN 3.5 [39]
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