Sophie Von Der Heyden

Maria Beger

Robert J Toonen

Lynne Van Herwerden

Marie Antonette Juinio-Meñez

Rachel Ravago-Gotanco

Cecile Fauvelot

Giacomo Bernardi

The application of genetics to marine management and conservation: examples from the Indo-Pacific

ABSTRACT.-Molecular tools and analyses have played pivotal roles in uncovering the processes and patterns of biodiversity in the Indian and Pacific oceans. However, integrating genetic results into management and conservation objectives has been challenging, with few examples that show practical applicability. This review aims to address some of the perceived barriers to an enhanced approach that integrates molecular data into management and conservation goals, by reviewing papers relevant to both conservation and fisheries management in the Indo-Pacific region, particularly with respect to phylogeography, connectivity, and species identification, as well as stock delineation, restoration of depleted wild stocks, mislabeled marine resources and "molecular forensics." We also highlight case studies from each of these areas that illustrate how molecular analyses are relevant to conservation and management in the Indo-Pacific, spanning a variety of vertebrate and invertebrate species. We discuss the application of genetic data to the design and evaluation of the effectiveness of marine protected area networks, stock delineation, and restoration and the usage of exclusion tests and parentage analyses for fisheries management. We conclude that there is a distinct need for increasing public awareness and ownership of genetically unique lineages and, ultimately, the increased inclusion of genetic research into management policy and conservation. Finally, we make a case for the importance of clear and effective communication for promoting public awareness, public ownership, and for achieving conservation goals within the region.

The Indo-Pacific, which encompasses large areas of the tropical Indian and Pacific oceans, houses an incredible array of marine biodiversity, with almost 4000 described species of fishes (Allen 2007), >700 tropical reef corals [START_REF] Connolly | Indo-Pacific biodiversity of coral reefs: deviations from a mid-domain model[END_REF], an undescribed diversity of invertebrates [START_REF] Bouchet | Assessing the magnitude of species richness in tropical marine environments: exceptionally high numbers of molluscs at a New Caledonia site[END_REF], Appletans et al. 2012), and is thus celebrated as a prominent marine biodiversity hotspot. Covering a large geographical area, it encompasses both developed and developing countries, which face vastly different pressures from global and climate change. Many of these pressures, including increasing sea-surface temperatures, ocean acidification, pollution, legal and illegal overfishing, and destructive fishing methods, have negative impacts on biodiversity. Thus, there is a clear need for improved conservation efforts in the region, which also allow sustainable exploitation and development of all Indo-Pacific nations.

Genetic approaches are one tool that can help inform the conservation and management objectives unique to marine systems. The past 20 yrs have seen a significant increase in the use of molecular tools addressing marine species evolution, biodiversity, and distributions. There is a wide array of studies, which at the broader scale include phylogeography (including comparative phylogeography), connectivity and migration, speciation and diversification, to more fine-scale studies that examine parentage analyses, population dynamics and demographic change, barcoding initiatives, and invasive species. Despite significant numbers of papers published, many state only the necessity of using genetic data for management and conservation; largely such data are not applied to real-life situations. This is primarily because of a lack of theoretical and practical frameworks to guide the use of genetic data [START_REF] Beger | Evolving coral reef conservation with genetic information[END_REF]. Other discrepancies arise from the differing approach and terminology in the disparate fields of conservation genetics, spatial conservation science, and fisheries management. Further, the multitude of genetic approaches, their interpretation, and follow-on research are often difficult for non-specialists to grasp; this challenge is addressed by Bowen et al. (2014).

Genetic research in the Indo-Pacific has been carried out on a diverse array of invertebrate and vertebrate species and has spanned the limits of the oceans, from southern Africa and the east African coast to the west coast of the United States and Australia. A brief web-based literature search carried out in Web of Science using the following search terms (Indo-Pacific and gene*, Indo-Pacific and fish* man*, Indo-Pacific and marine conservation, Indo-Pacific and pop* gen*) between January 2009 and April 2012 suggests that the most common research threads are phylogeography and population genetics, followed by phylogenetics and taxonomy and the development of new molecular markers (data not shown; see also [START_REF] Keyse | The scope of published population genetic data for Indo-Pacific marine fauna and future research opportunities in the region[END_REF]. There is wide taxonomic coverage, with corals and fishes being especially well represented. Some invertebrate taxa including crustaceans and echinoderms (particularly those of commercial interest) have been reasonably well studied, but in the context of conservation and management, many smaller invertebrate taxa such as molluscs, polychaete, and annelid worms and sponges are dramatically under represented.

This review aims to address some of the perceived barriers to an enhanced approach that integrates molecular data into management and conservation goals, by reviewing papers and highlighting case studies that apply a genetic approach to address issues around: (1) species identification and (2) spatial management within the Indo-Pacific region (Fig. 1). Specific case studies are selected to illustrate the application of genetic tools across a variety of conservation and management issues within each of these broad themes. Future applications of molecular tools in the region are then briefly discussed.

Species Identification

Identifying Cryptic Species and Hybridization Events to Better Characterize Biodiversity of the Indo-Pacific.-Species are cryptic when two or more reproductively isolated species are classified as one recognized species, because, most commonly, phenotypic characters that distinguish them are either absent or share a large degree of overlap. Accurate species inventories that are skewed by undescribed cryptic species are important for several reasons including precise biodiversity inventories, which are crucial for understanding ecological functions. Cryptic species also may play a large role in the management of exploited species. For example, the giant clam Tridacna squamosina (= T. costata in [START_REF] Richter | Collapse of a new living species of giant clam in the Red Sea[END_REF], see Online Appendices for species authorities) was shown to be genetically, ecologically, and morphologically distinct from Tridacna maxima, and T. squamosa, and at the same time to have been exploited for at least 125,000 yrs, with substantial overfishing reducing T. squamosina to <1% of stocks of giant clam in the Red Sea [START_REF] Richter | Collapse of a new living species of giant clam in the Red Sea[END_REF].

Tropical marine systems, such as those found in the Indo-Pacific, have been highlighted as potentially rich in cryptic species given that they are among the most species-rich habitats and as such should harbor greater numbers of cryptic and/or sibling species [START_REF] Bickford | Cryptic species as a window on diversity and conservation[END_REF]. For example, von der Heyden (2011) suggests that most cryptic fish species in southern Africa would belong to the western Indian Ocean fish fauna rather than the cool-temperate Atlantic.

Molecular tools have been at the forefront of identifying cryptic species. Interestingly, rather than suspecting that a single species may actually be composed of multiple species and then testing this hypothesis, most cryptic species discoveries are probably accidental. Barcoding initiatives and phylogeographic surveys especially seem to discover unknown biodiversity [START_REF] Barber | Estimating diversity of Indo-Pacific coral reef stomatopods through DNA barcoding of stomatopod larvae[END_REF][START_REF] Concepcion | An alternative to ITS, a hypervariable, single-copy nuclear intron in corals, and its use in detecting cryptic species within the octocoral genus Carijoa[END_REF][START_REF] Zemlak | DNA barcoding reveals overlooked marine fishes[END_REF][START_REF] Dibattista | Phylogeography of the bluelined surgeonfish, Acanthurus nigroris, reveals high connectivity and a cryptic endemic species in the Hawaiian Archipelago[END_REF][START_REF] Hubert | Cryptic diversity in Indo-Pacific coral-reef fishes revealed by DNA-barcoding provides new support to the Centre-of-Overlap hypothesis[END_REF], and such studies can afford an excellent view into the processes driving speciation in marine systems (von der Heyden et al. 2011).

In the understudied Indo-Pacific, numerous cryptic species have been described including fishes [START_REF] Colborn | The evolutionary enigma of bonefishes (Albula spp.): cryptic species and ancient separations in a globally distributed shorefish[END_REF][START_REF] Messmer | Phylogeography of colour polymorphism in the coral reef fish Pseudochromis fuscus, from Papua New Guinea and the Great Barrier Reef[END_REF][START_REF] Hubert | Cryptic diversity in Indo-Pacific coral-reef fishes revealed by DNA-barcoding provides new support to the Centre-of-Overlap hypothesis[END_REF], sponges [START_REF] Muricy | Diversity of Indo-Australian Plakortis (Demospongiae: Plakinidae), with description of four new species[END_REF], several groups of crustaceans (Barber andBoyce 2006, Poore andAndreakis 2011), molluscs [START_REF] Bouchet | Assessing the magnitude of species richness in tropical marine environments: exceptionally high numbers of molluscs at a New Caledonia site[END_REF][START_REF] Pola | Molecular data illuminate cryptic nudibranch species: the evolution of the Scyllaeidae (Nudibranchia: Dendronotina) with a revision of Notobryon[END_REF], echinoderms [START_REF] Landry | Recent speciation in the Indo-West Pacific: rapid evolution of gamete recognition and sperm morphology in cryptic species of sea urchin[END_REF][START_REF] Hart | Morphological and genetic variation indicate cryptic species within Lamarck's little sea star, Parvulastra (=Patiriella) exigua[END_REF][START_REF] Uthicke | Genetic barcoding of commercial Beche-de-mer species (Echinodermata: Holothuroidea)[END_REF], and jellyfishes (Dawson andMartin 2001, Holland et al. 2004). Uncovering cryptic species, either as sibling species or as species flocks, allows rare insights into the processes driving speciation in marine environments, especially where species cooccur without evidence of strong vicariant barriers (von der Heyden et al. 2011, Bowen et al. 2013). Such studies can have direct management consequences, such as the pending petition to list rare species of corals under the US Endangered Species Act, where the species status of many of those corals is in dispute (e.g., [START_REF] Forsman | Ecomorph or endangered coral? DNA and microstructure reveal Hawaiian species complexes: Montipora dilatata/flabellata/turgescens & M. patula/verrilli[END_REF]. Some of the strongest evidence for both cryptic species comes from coral reef fishes [START_REF] Rocha | Speciation in coral-reef fishes[END_REF], which display morphological variation that can be used as a baseline for genetic investigations. For example, in the genus Centropyge, mtDNA revealed deep geographic separation that is incongruent with species delineation, whereas nuclear analyses suggest ongoing gene flow between species [START_REF] Dibattista | Twisted sister species of pygmy angelfishes: discordance between taxonomy, coloration, and phylogenetics[END_REF]. It is often difficult to pinpoint the processes involved in speciation and the maintenance of species boundaries, but assortative mating and other selective forces have been suggested [START_REF] Palumbi | Genetic divergence, reproductive isolation, and marine speciation[END_REF][START_REF] Hellberg | Sympatric sea shells along the sea's shore: the geography of speciation in the marine gastropod Tegula[END_REF][START_REF] Mcmillan | Color pattern evolution, assortative mating, and genetic differentiation in brightly colored butterflyfishes (Chaetodontidae)[END_REF].

Hybridization is another complex process that skews our understanding of traditional species, although it may be somewhat easier to detect when parental species are morphologically distinct. Most hybrids in contact zones studied to date show at least some morphological intermediacy with both parent species, although this is not always apparent [START_REF] Pyle | A review of hybridisation in marine angelfishes (Perciformes: Pomacanthidae)[END_REF][START_REF] Mcmillan | Color pattern evolution, assortative mating, and genetic differentiation in brightly colored butterflyfishes (Chaetodontidae)[END_REF], van Herwerden et al. 2006[START_REF] Van Der Meer | Historic hybridisation and introgression between two iconic Australian anemonefish and contemporary patterns of population connectivity[END_REF][START_REF] Der Heyden | Evidence of hybridisation within the genus Chrysoblephus and conserved nuclear sequences of South African sparids (Teleostei: Sparidae)[END_REF]. Hybrids have been recorded in fishes, echinoderms, molluscs, and corals of the region [START_REF] Lessios | Hybridisation and introgression between Indo-Pacific species of Diadema[END_REF], van Herwerden et al. 2006[START_REF] Yaakub | Hybridisation in coral reef fishes: introgression and bi-directional gene exchange in Thalassoma (family Labridae)[END_REF][START_REF] Richards | Some rare Indo-Pacific coral species are probable hybrids[END_REF][START_REF] Hobbs | Marine hybrid hotspot at Indo-Pacific biogeographic border[END_REF][START_REF] Montanari | Reef fish hybridisation: lessons learnt from butterflyfishes (genus Chaetodon)[END_REF][START_REF] Ladner | Extensive sympatry, cryptic diversity and introgression throughout the geographic distribution of two coral species complexes[END_REF][START_REF] Van Der Meer | Historic hybridisation and introgression between two iconic Australian anemonefish and contemporary patterns of population connectivity[END_REF][START_REF] Zigler | Natural hybridisation in the sea urchin genus Pseudoboletia between species without apparent barriers to gamete recognition[END_REF]).

Hybridization appears to be common between species that underwent historical allopatry, i.e., for example during sea-level changes that created vicariant barriers, followed by subsequent contact of closely related sister lineages. Hybridization studies can contribute toward our understanding of historical processes involved in speciation events; however, they can also confound population genetic analyses (and therefore potentially also fisheries or management assessments) if hybrids prove problematic to separate (e.g., [START_REF] Addison | Colonization, dispersal, and hybridisation influence phylogeography of North Atlantic sea urchins (Strongylocentrotus droebachiensis)[END_REF]. For example coral trout, Plectropomus leopardus, is the primary commercial fishery on the Great Barrier Reef, where it has hybridized historically with a closely related congener, Plectropomus maculatus, and mtDNA does not differentiate them, although mtDNA distinguishes them elsewhere in their range [START_REF] Van Herwerden | Contrasting patterns of genetic structure in two species of the coral trout Plectropomus (Serranidae) from east and west Australia: introgressive hybridisation or ancestral polymorphisms[END_REF], also see [START_REF] Morgan | Detection of interspecies hybridisation in Chondrichthyes: hybrids and hybrid offspring between Australian Carcharhinus tilstoni and common C. limbatus blacktip shark found in an Australian fishery[END_REF].

There are several regions within the Indo-Pacific that have been highlighted for potential interest in promoting hybridization in marine species. For example, the eastern Indian Ocean at the Christmas and Cocos islands has been suggested as a marine hybrid hotspot in fishes [START_REF] Hobbs | Marine hybrid hotspot at Indo-Pacific biogeographic border[END_REF]). The Indo-Pacific barrier also influenced the divergence of marine species during glacial cycles, with subsequent overlap of sister lineages or species [START_REF] Gaither | High connectivity in the Deepwater Snapper Pristipomoides filamentosus (Lutjanidae) across the Indo-Pacific with isolation of the Hawaiian Archipelago[END_REF]. Other examples of processes driving hybridization between species include those where hybridization appears to be driven by unequal parent ratios, i.e., if one parent species is rare, mating with conspecific hybrids increases [START_REF] Marie | Hybridisation of reef fishes at the Indo-Pacific biogeographic barrier: a case study[END_REF][START_REF] Hobbs | Marine hybrid hotspot at Indo-Pacific biogeographic border[END_REF][START_REF] Burford | Analysis of individual year-classes of a marine fish reveals little evidence of first-generation hybrids between cryptic species in sympatric regions[END_REF][START_REF] Montanari | Reef fish hybridisation: lessons learnt from butterflyfishes (genus Chaetodon)[END_REF][START_REF] Der Heyden | Evidence of hybridisation within the genus Chrysoblephus and conserved nuclear sequences of South African sparids (Teleostei: Sparidae)[END_REF]. Human or ecological disturbance may also increase the rate of hybridization in fishes [START_REF] Van Herwerden | Phylogenetic and evolutionary perspectives of the Indo-Pacific grouper Plectropomus species on the Great Barrier Reef, Australia[END_REF][START_REF] Mullen | Hybridisation among Caribbean damselfish species correlates with habitat degradation[END_REF], although whether this also holds for invertebrate species in the Indo-Pacific region remains a large research gap (but see [START_REF] Uthicke | Natural hybridisation does not dissolve species boundaries in commercially important sea cucumbers[END_REF]. Further, whether rates of hybridization in marine species will increase with global change scenarios remains to be tested, but this has been suggested for at least one pair of hybridizing shark species [START_REF] Morgan | Detection of interspecies hybridisation in Chondrichthyes: hybrids and hybrid offspring between Australian Carcharhinus tilstoni and common C. limbatus blacktip shark found in an Australian fishery[END_REF].

Detection of Invasive Species and Reconstruction of Invasion

Histories.-Invasive species have played a large role in changing global marine systems, in some cases drastically altering the biological communities of invaded areas [START_REF] Blum | The nonnative solitary ascidian Ciona intestinalis (L.) depresses species richness[END_REF][START_REF] Robinson | Changes in South African rocky intertidal invertebrate community structure associated with the invasion of the mussel Mytilus galloprovincialis[END_REF][START_REF] Banks | Genetic structure of a recent climate change-driven range extension[END_REF], and are recognized as a pressing global concern [START_REF] Sutherland | A horizon scan of global conservation issues for 2010[END_REF]). The Indo-Pacific is no exception, with a number of non-indigenous species (NIS) recorded throughout the region. Monitoring of invasive species is critical for understanding the biological and economic consequences of invasions, although the extent of invasive species is unknown in many developing countries that do not necessarily have the resources and capacity to track marine invasions. Molecular tools are able to provide an additional dimension in our understanding of the dynamics of marine species invasions, such as resolving the genetic relationships of invasive species with native species, monitoring potential hybridization of invasive and native species, and unraveling the geographic provenance of native populations. For example, for seaweeds in southern Africa [START_REF] Bolton | Molecular evidence for three separate cryptic introductions of the red seaweed Asparagopsis (Bonnemaisoniales, Rhodophyta) in South Africa[END_REF], fishes [START_REF] Gaither | Identifying the perfect invader: population structure in the native range predicts the spread of introduced marine species[END_REF], and jellyfishes in the Hawaiian Islands [START_REF] Holland | Global phylogeography of Cassiopea (Scyphozoa : Rhizostomeae): molecular evidence for cryptic species and multiple invasions of the Hawaiian Islands[END_REF], species initially thought to be native have been exposed as being invasive, as morphological characteristics alone were not able to distinguish clearly between native and NIS.

Phylogeography has been successfully used to determine the likely founder populations or to track range expansions of marine NIS in the Indo-Pacific region. For the highly invasive green crab, Carcinus maenas, it was shown that South Africa experienced multiple invasions from the North Atlantic, with at least some individuals originating from northern Europe [START_REF] Darling | Genetic patterns across multiple introductions of the globally invasive crab genus Carcinus[END_REF]. For the invasive barnacle, Balanus glandula, which has outcompeted southern African Chthamalus dentatus, the invasive population could be traced back to the coast of Oregon [START_REF] Simon-Blecher | Balanus glandula: from north-west America to the west coast of South Africa[END_REF]). To the contrary, four invasive species of ascidians appear to have experienced decades of anthropogenically mediated transport so that no genetic signal of potential native populations can be distinguished [START_REF] Rius | Range expansions across ecoregions: interactions of climate change, physiology and genetic diversity[END_REF]. Alternative examples also exist where invasive species thought to be alien were shown to be native rather than introduced. A good example is the snowflake coral, Carijoa, which was thought to be introduced into the Hawaiian archipelago from the Caribbean, yet molecular analyses revealed that Carioja was native to the Indo-Pacific, rather than introduced from the Caribbean-Atlantic [START_REF] Concepcion | Resolving natural ranges and marine invasions in a globally distributed octocoral (genus Carioja)[END_REF]. This clearly highlights the need for not only genetic monitoring of invasive species, but that molecular systematics remain highly important in evaluating the status of invasive and NIS.

Molecular tools become invaluable when examining species without diagnostic morphological features. For example, endosymbiotic Symbiodinium dinoflagellates could potentially be dispersed to and associate with new coral hosts via natural and anthropogenic means. [START_REF] Stat | Vectored introductions of marine endosymbiotic dinoflagellates into Hawaii[END_REF] detected an unusual Symbiodinium lineage A1 in Acropora cythera from the Northwestern Hawaiian Islands and suggested that this may have been transported into Hawaii by an NIS jellyfish vector. Even larger animals such as seahorses have proven confusing; Hippocampus kuda in Hawaii is thought to be an introduced species through the aquarium trade. However, molecular, as well as careful morphological analyses show that seahorses in Hawaii are genetically and morphologically distinct from H. kuda [START_REF] Szabó | On the status of the Hawaiian seahorses Hippocampus hilonis, H. histrix and H. fisheri (Syngnathidae)[END_REF]) and that they are not introduced. Both studies highlighted above support the need for large-scale geographic sampling for putative NIS to prevent "false positives" and a potential misuse of funds for monitoring so-called NIS, which turn out to be indigenous. Such extensive sampling will of course be more demanding for smaller-sized species or those with extensive global distributions and is emphasized in a paper by [START_REF] Bolton | Jellyfish on the rocks: bioinvasion threat of the international trade in aquarium live rock[END_REF], which highlights the dangers of trade in live aquatic species that may disperse invasive species such as jellyfishes (that may be morphologically indistinguishable) into new environments.

Market-based Surveys, Mislabeling of Marine Resources, and Molecular Forensics.-Globally, fisheries resources are under pressure, with the majority of fish stocks fully exploited, overexploited, or depleted [START_REF] Anticamara | Global fishing effort (1950-2010): trends, gaps and implications[END_REF], including some of those in the Indo-Pacific [START_REF] Pauly | Fishing down marine food webs[END_REF]. Seafood fraud, whether through the intentional mislabeling of species or accidentally misidentifying or misnaming catches, is under-reported globally, with most examples from North America [START_REF] Marko | Mislabelling of a depleted reef fish[END_REF][START_REF] Wong | DNA barcoding detects market substitution in North American seafood[END_REF], Hanner et al. 2011[START_REF] Cline | Marketplace substitution of Atlantic salmon for Pacific salmon in Washington State detected by DNA barcoding[END_REF] and Europe (Miller andMariani 2010, Garcia-Vasquez et al. 2011). Fraud in the seafood market most commonly takes place when low-value or more readily available species are substituted for expensive species or those that are rare and in highdemand, including protected species such as whales [START_REF] Baker | Molecular genetic identification of whale and dolphin products from commercial markets in Korea and Japan[END_REF][START_REF] Jacquet | Trade secrets: renaming and mislabelling of seafood[END_REF], Marko et al. 2011[START_REF] Cline | Marketplace substitution of Atlantic salmon for Pacific salmon in Washington State detected by DNA barcoding[END_REF]). This especially is prevalent in processed products, where diagnostic characters are removed. Molecular tools, primarily barcoding approaches, are therefore most commonly used to identify mislabeled seafood products in the market place (Ogden 2008, Rasmussen and[START_REF] Rasmussen | DNA-based methods for the identification of commercial fish and seafood species[END_REF]. Within the Indo-Pacific, however, levels of seafood mislabeling and fraud remain almost unknown as only a few studies have been carried out in the region.

In Japan, developments of a reliable PCR-RFLP method has allowed for the discrimination of locally caught chub mackerel, Scomber japonicas, and imported Atlantic mackerel, Scomber scombrus [START_REF] Aranishi | Rapid PCR-RFLP method for discrimination of imported and domestic mackerel[END_REF], although no market-related validation was carried out. Likewise, whale meat being sold or served in Korea and Japan was "spot-checked" using mtDNA sequences and shown to represent at least six species of whale and some dolphin species [START_REF] Baker | Molecular genetic identification of whale and dolphin products from commercial markets in Korea and Japan[END_REF]. In South Africa, >50% of fish fillets tested (comprising four different species) were mislabeled, with kob, Argyrosomus spp., having a substitution rate of >80% (von der Heyden et al. 2010). Further, over half of the fishes that were found to be substitutes were not recorded from southern African waters, suggesting large-scale import and renaming of marine species from elsewhere, including Australia [START_REF] Der Heyden | Misleading the masses: detection of mislabelled and substituted frozen fish products in South Africa[END_REF]. Protocols also exist to identify shark fins and assign them to species [START_REF] Shivji | Genetic identification of shark body parts for conservation and trade monitoring[END_REF] and the largest shark fin market in the world, Hong Kong, has been monitored using genetic tools to identify the most heavily traded species [START_REF] Clarke | Identification of shark species composition and proportion in the Hong Kong shark fin market based on molecular genetics and trade records[END_REF]. This application of genetics is vital to conservation and fisheries management, because the largely unregulated trade in shark fins is one of the most serious threats to shark populations worldwide. [START_REF] Magnussen | Genetic tracking of basking shark products in international trade[END_REF] also developed a species-specific genetic assay to monitor trade in basking shark (Cetorhinus maximus) products, as this Appendix II CITES listed species is particularly sensitive to exploitation due to premium prices achieved for its products.

Barcoding initiatives [START_REF] Lemer | Cytochrome b barcoding, molecular systematics and geographic differentiation in rabbitfishes (Siganidae)[END_REF][START_REF] Wörheide | DNA taxonomy of sponges-progress and perspectives[END_REF][START_REF] Plaisance | Reef-associated crustacean fauna: biodiversity estimates using semi-quantitative sampling and DNA barcoding[END_REF][START_REF] Uthicke | Genetic barcoding of commercial Beche-de-mer species (Echinodermata: Holothuroidea)[END_REF][START_REF] Hubert | Cryptic diversity in Indo-Pacific coral-reef fishes revealed by DNA-barcoding provides new support to the Centre-of-Overlap hypothesis[END_REF]) within the Indo-Pacific region should therefore be expanded to not only assess species substitutions, but also provenance of marine species. For example, fish marketed as "South African" kingklip, Genypterus capensis, was found to be substituted by pink ling, Genypterus blacodes, caught in New Zealand waters [START_REF] Der Heyden | Misleading the masses: detection of mislabelled and substituted frozen fish products in South Africa[END_REF]. Further, the use of barcoding to discriminate between populations of swordfish, Xiphias gladius, suggests that the COI gene is sensitive enough to separate fishes caught in the Atlantic and Indian oceans, as well as the Mediterranean [START_REF] Pappalardo | Geographically widespread swordfish barcode stock identification: a case study of its application[END_REF], although the study did not reveal any substitution with fishes caught in the Indian Ocean. Barcoding has also been successfully applied in monitoring the shark fishery in Madagascar [START_REF] Doukakis | Applying genetic techniques to study remote shark fisheries in northeastern Madagascar[END_REF] determining the most commonly marketed species.

For molecular methods to be useful in determining levels of seafood fraud and mislabeling in the Indo-Pacific, overall monitoring of fishing activities and reporting of catches (at local and regional scales) needs to be drastically improved. Further, resolving cryptic species and describing unknown biodiversity will also greatly aid in molecular techniques being applied to tracing products within the seafood chain and combating mislabeling. Ultimately, an approach that allows individuals to be assigned to populations [using for example Single Nucleotide Polymorphisms (SNPs) such as used by the FishPopTrace Consortium (2012)], rather than simply species, will ensure more effective policing of Indo-Pacific marine resources, although this has to date not been applied widely and remains a challenge globally.

Spatial Planning and Restoration

Spatial Management of Wild Marine Species within the Indo-Pacific.-Knowledge of the scale and direction of connectivity among sites is pertinent to effective marine conservation [START_REF] Almany | Connectivity, biodiversity conservation and the design of marine reserve networks for coral reefs[END_REF], McCook et al. 2009[START_REF] Beger | Incorporating functional ecological connectivity into spatial decision making for conservation[END_REF][START_REF] Kininmonth | Dispersal connectivity and reserve selection for marine conservation[END_REF][START_REF] Olds | Habitat connectivity improves reserve performance[END_REF]) and fisheries management (e.g., [START_REF] Leis | Estimating connectivity in marine fish populations: what works best?[END_REF]. Particularly when combining genetic information with biophysical models [START_REF] Kininmonth | Determining the community structure of the coral Seriatopora hystrix from hydrodynamic and genetic networks[END_REF][START_REF] White | Ocean currents help explain population genetic structure[END_REF][START_REF] Kool | Connectivity and the development of population genetic structure in Indo-West Pacific coral reef communities[END_REF][START_REF] Foster | Connectivity of Caribbean coral populations: complementary insights from empirical and modelled gene flow[END_REF][START_REF] Treml | Marine population connectivity identifies ecological neighbors for conservation planning in the Coral Triangle[END_REF], genetic data can underpin the delineation of management areas by identifying consistently located breaks [START_REF] Carpenter | Comparative phylogeography of the Coral Triangle and implications for marine management[END_REF], Toonen et al. 2011;Fig. 1). For conservation or fisheries target species, genetic tools can assist with identifying and protecting genetically isolated populations [START_REF] Palsbøll | Identification of management units using population genetic data[END_REF] or cryptic species (Griffiths et al. 2010b) from overfishing and other threats. In the context of fisheries management, the identification of subpopulations from seemingly uniform stocks is also important, affecting spatial structuring of management (Table 1). For example, although deep-water snapper, Pristipomoides filamentosus, stocks are highly connected across the Indo-Pacific, the Hawaiian Archipelago supports an isolated subpopulation requiring separate fisheries management (Gaither et al. 2011). Similarly, a northern Australian cryptic species of the blue swimmer crab, Portunus pelagicus, discovered in an allozyme study highlighted the need for differential management measures [START_REF] Bryars | An allozyme study of the blue swimmer crab, Portunus pelagicus (Crustacea : Portunidae), in Australia: stock delineation in southern Australia and evidence for a cryptic species in northern waters[END_REF]. Genetic structure relevant to fisheries management can sometimes be observed at inter-regional and larger spatial scales, but not within regions or locally such as in coral trout, Plectropomus leopardus (Table 1). Where highly structured populations relate to localized circulation patterns, such as for fisheries species in the South Californian Bight, genetic analysis can identify source sites to protect for maintaining sustainable fisheries (Table 1; [START_REF] Selkoe | Effects of ephemeral circulation on recruitment and connectivity of nearshore fish populations spanning Southern and Baja California[END_REF][START_REF] White | Ocean currents help explain population genetic structure[END_REF]. Similarly, the identification and incorporation of such population source-sink dynamics is among the last remaining frontiers in marine spatial planning [START_REF] Crowder | Source-sink population dynamics and the problem of siting marine reserves[END_REF][START_REF] Bode | Larval dispersal reveals regional sources and sinks in the Great Barrier Reef[END_REF][START_REF] López-Duarte | What controls connectivity? An empirical, multi-species approach[END_REF]).

In the context of marine spatial planning for biodiversity conservation, the delineation of genetically differentiated subpopulations that are treated as management units aids the spatial structuring of conservation measures that aim to represent all facets of biodiversity. Several examples utilize a variety of genetic approaches to allocate organisms to management units, such as basing the grouping of sites for management on a measure of connectivity among sites [START_REF] Kininmonth | Determining the community structure of the coral Seriatopora hystrix from hydrodynamic and genetic networks[END_REF], Toonen et al. 2011), evolutionary significant units (Barber et al. 2006), phylogeography [START_REF] Reece | Comparative phylogeography of four Indo-Pacific moray eel species (Muraenidae) reveals comparable ocean-wide genetic connectivity despite five-fold differences in available adult habitat[END_REF], and evolutionary processes [START_REF] Crandall | Expansion dating: calibrating molecular clocks in marine species from expansions on the Sunda shelf following the last glacial maximum[END_REF]. To include measures of connectivity in spatial planning, genetically differentiated units, akin to management units, were determined in a model that combined F ST values between pairs of sites with connectivity values derived from a biophysical model and biogeographic measures [START_REF] Kininmonth | Determining the community structure of the coral Seriatopora hystrix from hydrodynamic and genetic networks[END_REF]. Comparative phylogeography across several taxa in the Philippines [START_REF] Carpenter | Comparative phylogeography of the Coral Triangle and implications for marine management[END_REF] and Hawaii [START_REF] Toonen | Defining boundaries for applying ecosystem-based management: a multispecies case study of marine connectivity across the Hawaiian Archipelago[END_REF]) was used to highlight potential management units for spatial planning aiming to preserve core evolutionary processes and histories. Marine phylogeography is also informative for identifying management units, evolutionary significant units, cryptic species, and endemism [START_REF] Rocha | Phylogeography and the conservation of coral reef fishes[END_REF]). Isolated evolutionary significant areas are defined when spatially and genetically differentiated patterns in local populations of multiple species are repeatedly identified at specific locations; for example, various coral reef fish species at the eastern edge of the west Pacific-Hawaii, Marquesas, and French Polynesia [e.g., the parrotfishes Chlorurus sordidus [START_REF] Bay | High genetic diversities and complex genetic structure in an Indo-Pacific tropical reef fish (Chlorurus sordidus): evidence of an unstable evolutionary past?[END_REF]) and Scarus psittacus, [START_REF] Winters | Phylogeography of the Indo-Pacific parrotfish Scarus psittacus: isolation generates distinctive peripheral populations in two oceans[END_REF]; the snappers Lutjanus fulvus and Lutjanus kasmira [START_REF] Gaither | Genetic evaluation of marine biogeographical barriers: perspectives from two widespread Indo-Pacific snappers (Lutjanus kasmira and Lutjanus fulvus)[END_REF], the boring clam, Tridacna crocea, [START_REF] Deboer | Phylogeography and limited genetic connectivity in the endangered boring giant clam across the Coral Triangle[END_REF]Fig. 1)]. Populations at such isolated locations should be managed independently to preserve genetic diversity. Genetic approaches are also highly relevant to benchmark protected area and management success. For example, DNA genetic fingerprinting allowed the monitoring of stock recovery for overfished holothurians on the Great Barrier Reef in combination with repeated large scale surveys to identify stock status (Uthicke et al. 2004). On smaller spatial and time scales, parentage analysis has been used to establish patterns of self-recruitment in anemone fishes, Amphiprion polymnus [START_REF] Jones | Coral reef fish larvae settle close to home[END_REF], gobies, Elacatinus evelynae [START_REF] Taylor | Genetic evidence for local retention of pelagic larvae in a Caribbean reef fish[END_REF], butterfly fishes, Chaetodon vagabundus [START_REF] Almany | Local replenishment of coral reef fish populations in a marine reserve[END_REF], yellow tang, Zebrasoma flavescens, damselfish, Stegastes partitus (Christie et al. 2010a,b) and striped snappers, Lutjanus carponotatus [START_REF] Harrison | Larval export from marine reserves and the recruitment benefit for fish and fisheries[END_REF], helping to quantify recruitment patterns in and out of reserves. Similarly, the rates of dispersal from a parent site to other suitable habitats in the region were identified for an anemone fish and a butterfly fish in Kimbe Bay, Papua New Guinea, with parentage analysis as the most robust tool for quantifying connectivity under high gene flow scenarios [START_REF] Saenz-Agudelo | Estimating connectivity in marine populations: an empirical evaluation of assignment tests and parentage analysis under different gene flow scenarios[END_REF].

Genotype assignments in conjunction with mtDNA sequence analyses have also been used successfully to identify insular, self-seeding local populations of inshore fishes displaying low gene flow in tropical northern Australia, e.g., the polynemid fishes Eleutheronema tetradactylum [START_REF] Horne | Limited ecological population connectivity suggests low demands on self-recruitment in a tropical inshore marine fish (Eleutheronema tetradactylum: Polynemidae)[END_REF]) and Polydactylus macrochir [START_REF] Horne | Searching for common threads in threadfins: phylogeography of Australian polynemids in space and time[END_REF]. Regardless of the method, estimates of connectivity and dispersal distances determined with genetic tools are important for choosing the most effective size of marine reserves to protect recruitment processes, which has been applied to corals [START_REF] Van Oppen | Genetic traces of recent long-distance dispersal in a predominantly self-recruiting coral[END_REF][START_REF] Underwood | Ecologically relevant dispersal of corals on isolated reefs: implications for managing resilience[END_REF]) and fishes [START_REF] Almany | Local replenishment of coral reef fish populations in a marine reserve[END_REF][START_REF] Buston | Probability of successful larval dispersal declines fivefold over 1 km in a coral reef fish[END_REF][START_REF] Harrison | Larval export from marine reserves and the recruitment benefit for fish and fisheries[END_REF]. A number of caveats remain when calculating dispersal distances from genetic data [START_REF] Palumbi | Population genetics, demographic connectivity, and the design of marine reserves[END_REF][START_REF] Hellberg | Gene flow and isolation among populations of marine animals[END_REF]. These include inferring a lack of dispersal as the cause of differentiation [START_REF] Hart | Next-generation studies of mating system evolution[END_REF][START_REF] Hart | It's about time: divergence, demography, and the evolution of developmental modes in marine invertebrates[END_REF][START_REF] Marko | The complex analytical landscape of gene flow inference[END_REF], the influence of pelagic larval duration (PLD) on dispersal ability [START_REF] Weersing | Population genetics, larval dispersal, and demographic connectivity in marine systems[END_REF][START_REF] Selkoe | Marine connectivity: a new look at pelagic larval duration and genetic metrics of dispersal[END_REF][START_REF] Riginos | Effects of geography and life history traits on genetic differentiation in benthic marine fishes[END_REF][START_REF] Faurby | Theoretical limits to the correlation between pelagic larval duration anfd population genetic structure[END_REF]) and realized dispersal on population genetic structure and gene flow.

Further, other issues that are often not addressed pertain to: (1) differences in stock structure and genetic connectivity when measured using markers assumed to be either neutral (due to genetic drift alone) or under selection (adaptive; but see Bowen et al. 2014); conservation genomics has set the stage to address this, so we expect to see such studies emerging in the coming years. ( 2) Differences in genetic structure may be detected due to rarely examined temporal variation, but where it has been examined, it is unclear how this affects larger-scale structure. This may also depend on the biological characteristics of the study species (e.g., [START_REF] Bjorndal | Annual variation in source contributions to a mixed stock: implications for quantifying connectivity[END_REF].

Genetic Studies of Stock Enhancement and Restoration in the Indo-Pacific.-Release of cultured juveniles has been undertaken to rebuild or enhance wild stocks (e.g., [START_REF] Bell | Restocking and stock enhancement of marine invertebrate fisheries[END_REF]. Aside from animal health considerations, there are concerns about negative impacts on the wild gene pool due to the relocation of individuals and/or the release of hatchery-produced individuals into the natural environment [START_REF] Blankenship | A responsible approach to marine stock enhancement[END_REF]Leber 1995, Tringali and[START_REF] Tringali | Risk to genetic effective population size should be an important consideration in fish stock-enhancement programs[END_REF]. In theory, the loss of genetic diversity through introgression and interbreeding of cultured and wild stocks may result in decreased fecundity, slowed growth, and high mortality [START_REF] Uthicke | Preservation of genetic diversity in restocking of the sea cucumber Holothuria scabra investigated by allozyme electrophoresis[END_REF]); these effects can occur within a single generation [START_REF] Christie | Genetic adaptation to captivity can occur in a single generation[END_REF]. Coupled with environmental stress, a decrease in the fitness of a species may eventually result in extinction. It has also been suggested that maintaining high genetic diversity is important in strengthening the resilience of the wild population to changes in environmental conditions [START_REF] Gamfeldt | Increasing intraspecific diversity enhances settling success in a marine invertebrate[END_REF]. In relation to these potential genetic effects, genetic characteristics of the natural populations should be considered before any stock restoration and stock enhancement initiative [START_REF] Blankenship | A responsible approach to marine stock enhancement[END_REF]. As a precautionary measure, it is recommended that a large number of broodstock should be taken directly from the population to be enhanced and in cases where broodstock are no longer available from the area to be enhanced, the most genetically-similar populations should be used [START_REF] Ward | The importance of identifying spatial population structure in restocking and stock enhancement programmes[END_REF]). However, this would only be tenable if baseline population genetic information is available.

Few stock enhancement or restocking efforts in the Indo-Pacific have taken genetics into consideration. In Thailand, genetic differences of the giant clam, Tridacna squamosa, and the abalone, Haliotis asinina, from the Andaman Sea and Thailand Gulf have been investigated. Both species are cultured and commercially important and have potential for restocking. Isozyme analysis of the giant clam T. squamosa revealed significant differences in genotypic and allelic frequencies between populations from the two regions [START_REF] Kittiwattanawong | High genetic divergence of Tridacna squamosa living at the west and the east coasts of Thailand[END_REF]. Further, Random Amplification of Polymorphic DNA (RAPD) and microsatellite markers revealed high population genetic differences in founder populations of cultured Haliotis asinina [START_REF] Tang | Genetic heterogeneity of the tropical abalone (Haliotis asinina) revealed by RAPD and microsatellite analyses[END_REF]. Both studies suggested that stocks from these sites should be managed separately to maintain the unique genetic characteristics of the regions for fishery conservation and sustainable aquaculture. Restocking without considering wild population structure can have potentially disastrous effects on local populations. For example, the intentional stocking of the Japanese shortneck clam from China and the Korean peninsula to enhance depleted Japanese populations backfired when it turned out to be a cryptic species that is now rapidly hybridizing with the native population such that the hybrid proportion now exceeds 50% [START_REF] Kitada | Molecular and morphological evidence of hybridisation between native Ruditapes philippinarum and the introduced Ruditapes form in Japan[END_REF].

Commercial culture of sandfish, Holothuria scabra, has spurred interest in sea ranching and other culture-based initiatives in the Indo-West Pacific [START_REF] Eriksson | Sea cucumber aquaculture in the western Indian Ocean: challenges for sustainable livelihood and stock improvement[END_REF][START_REF] Purcell | Principles and science of stocking marine areas with sea cucumbers[END_REF]. Sandfish is among the most expensive tropical sea cucumber species and has thus been heavily exploited across the region (e.g., [START_REF] Heinen | Take the sea cucumber[END_REF][START_REF] Hasan | Destruction of a Holothuria scabra population by overfishing at Abu Rhamada Island in the Red Sea[END_REF][START_REF] Ahmed | The status of commercial sea cucumbers from Egypt's northern Red Sea Coast[END_REF]. Genetic surveys using allozymes on H. scabra populations from Indonesia, Torres Strait, in northeast Australia, Solomon Islands, and New Caledonia detected significant genetic structure among many populations, which indicate restricted gene flow despite the high dispersal potential of the larvae (Uthicke and Benzie 2001, [START_REF] Uthicke | Preservation of genetic diversity in restocking of the sea cucumber Holothuria scabra investigated by allozyme electrophoresis[END_REF]). Based on these results, it was recommended that management of H. scabra should be implemented at local scales and establishment of local marine reserves is also necessary [START_REF] Uthicke | Restricted gene flow between Holothuria scabra (Echinodermata: Holothuroidea) populations along the northeast coast of Australia and the Solomon Islands[END_REF]. Both studies also emphasized the importance of using locally-sourced spawners for hatchery production in relation to stock enhancement interventions. In this case, it is imperative to apply precautionary approaches to conserving genetic diversity in scaling-up culturebased management efforts over the entire range of this species.

Fine-scale Patterns of Local Recruitment within the Indo-Pacific.-Understanding recruitment dynamics of marine organisms is a key factor both at the theoretical level-as it links demographic, resilience, and succession issues-and at applied levels, since it informs managers about the effectiveness of the placement and distance between fragmented marine reserves. Parentage analyses are able to provide fine-scale resolution on recruitment dynamics in marine systems. Similar to forensic methods of paternity analyses, parentage analyses require a reasonable proportion of the adult population and many offspring to be genotyped using a large panel of microsatellite markers and SNPs (single nucleotide polymorphisms). Assignment tests are then carried out to link offspring and potential parents based on a likelihood estimate. The likelihood estimate of parentage association depends on the number of loci, their allelic diversity, and the distribution of alleles in the adult population. To use the parentage assignment method, one needs to find a species amenable to this type of analysis, where the species is not very abundant, and both adults and recruits are easy to find and sample. In a study carried out in Kimbe Bay, Papua New Guinea, anemonefish, Amphiprion polymnus, presented the perfect model system [START_REF] Jones | Coral reef fish larvae settle close to home[END_REF]) since they live exclusively in easy to find anemones that have low abundance, thus ensuring that most parents in a surveyed area are sampled. Results showed extreme local patterns of recruitment dynamics, and confirmed the early, coarser estimates of self-recruitment, based on tagged otoliths of the recruits [START_REF] Almany | Local replenishment of coral reef fish populations in a marine reserve[END_REF], Jones et al. 2005[START_REF] Saenz-Agudelo | Estimating connectivity in marine populations: an empirical evaluation of assignment tests and parentage analysis under different gene flow scenarios[END_REF], 2011). Interestingly, it was possible to directly compare the relative power of the genetic and otolith microchemistry approaches since specific individuals were both genotyped and used for otolith analyses. While otolith microchemistry provides a unique understanding of the type of chemical environment (water mass) larvae encounter each day before recruitment, genetic approaches were found to be more powerful in identifying the fraction of self-recruitment in the population [START_REF] Berumen | Persistence of self-recruitment and patterns of larval connectivity in a marine protected area network[END_REF], although some studies suggest otherwise (Lopez-Duarte et al. 2012). Besides simple recruitment assessment, precise patterns could be identified, such as the relative contribution of females of different sizes, where larger females were shown to disproportionally contribute to the self-recruiting population [START_REF] Beldade | Larger female fish contribute disproportionately more to self-replenishment[END_REF].

For parentage analysis studies to be broadly relevant, more species needed to be examined. The study of anemonefish was therefore expanded to one species of butterflyfish, Chaetodon vagabundus, in Kimbe Bay, Papua New Guinea [START_REF] Berumen | Persistence of self-recruitment and patterns of larval connectivity in a marine protected area network[END_REF], as well as yellow tangs and Caribbean damsels (Christie et al. 2010a,b). While these species present considerable challenges compared to the simpler anemonefish system, each was able to assign offspring to specific parents, proving that the general method of parentage analysis could be expanded to other reef fish species with more typical pelagic larvae. More importantly, some showed a direct application of parentage analysis approaches to conservation by linking their results to the efficiency and design of marine reserves (Berumen et al. 2012). Most parentage studies have used fishes, although an expansion of using parentage and kinship relationships comes from a recent study on California spiny lobsters, Panulirus interruptus, which require 240-330 d of pelagic development prior to settlement. Despite high gene flow and extensive mixing, [START_REF] Iacchei | Combined analyses of kinship and F ST suggest potential drivers of chaotic genetic patchiness in high gene-flow populations[END_REF] showed specific populations, associated with upwelling fronts, contained significantly more kin than expected and were thus significantly differentiated from other sites throughout the species range.

The design of marine protected areas (MPAs) has generally been based on a number of considerations, often not relevant to the biology of the species that they are supposed to protect [START_REF] Sala | The structure of Mediterranean rocky reef ecosystems across environmental and human gradients, and conservation implications[END_REF]. The application of parentage and kinship analyses to our understanding of connectivity among reefs is still in its infancy, yet it has yielded some very promising results that are likely to inform managers as to the proper design of future MPAs. New studies have now used parentage or kinship analyses to understand connectivity between reefs using species that are commercially important and highly relevant to fisheries managers. Specifically, bar-cheeked coral trout, Plectropomus maculatus, and Spanish flag snapper, Lutjanus carponotatus, from the Keppel Islands on the Great Barrier Reef, Australia, were used to identify the levels of connectivity between MPAs within the area, the levels of self-recruitment within MPAs and the levels of recruit spillover from MPAs into surrounding reefs open to fishing [START_REF] Harrison | Larval export from marine reserves and the recruitment benefit for fish and fisheries[END_REF]. This indicated how a network of MPAs may effectively sustain itself and areas open to fishing. Such studies have fueled a paradigm shift over the past decade, from the general idea of open populations to the discovery that self-recruitment may be far more prevalent than once imagined. We are still clearly at the beginning of applying the method. Indeed, microsatellites have inherent technical limitations and a comparison of the effectiveness of microsatellite and SNPs in parentage analysis favored the latter [START_REF] Hauser | An empirical comparison of SNPs and microsatellites for parentage and kinship assignment in a wild sockeye salmon (Oncorhynchus nerka) population[END_REF]. It is likely that next generation sequencing (NGS) approaches that rapidly identify and concurrently screen large numbers of SNPs across many individuals will be the method of choice for parentage analyses in non-model organisms in the near future [START_REF] Allendorf | Genomics and the future of conservation genetics[END_REF].

Management Cognizant of Changing Sex, Stock Structuring, and Group Spawning.-One difficulty in generalized management decisions stems from the unique life histories of species targeted for harvest in the Indo-Pacific, including sequential hermaphroditism and group spawning. Approximately 900 species of fish form the primary targets of artisanal and commercial fisheries in the Indo-Pacific. Targeted families with species which are sequential hermaphrodites include wrasses (Labridae), parrotfishes (Scaridae), emperors (Lethrinidae), and to some extent groupers (Serranidae) [START_REF] Nelson | Fishes of the World[END_REF]. When using genetic tools to assess populations, one needs to take into consideration this fundamental life history trait, as it may result in confusing genetic signatures that could be misinterpreted otherwise. For example, a classic interpretation of differences in the genetic results for male and female individuals is a difference in dispersal between genders, where one gender shows site fidelity and the other does not. For sequential hermaphrodites, such a trait, which is essential to proper management, would be masked in a genetic study where life history is not taken into account. Iconic species such as the humphead wrasse, Cheilinus undulatus, and the bumphead parrotfish, Bolbometopon muricatum, have been driven locally extinct in many places [START_REF] Bellwood | Limited functional redundancy in high diversity systems: resilience and ecosystem function on coral reefs[END_REF][START_REF] Sadovy | The humphead wrasse, Cheilinus undulatus: synopsis of a threatened and poorly known giant coral reef fish[END_REF][START_REF] Dulvy | Using informal knowledge to infer human-induced rarity of a conspicuous reef fish[END_REF]). In the past, management has generally recommended smaller size limit to catches, although it is now moving toward slot approaches where both size and quantities are taken into account. In the case of sequential hermaphrodites, where individuals change sex over the course of their lifetime, with younger individuals being one gender (female for protogynous hermaphrodites, male for protandrous hermaphrodites) and older individuals being the other gender, removing larger individuals results in a number of substantial population consequences. The natural disappearance of the largest individual triggers the sex change of the next largest individual [START_REF] Warner | Social control of sex change in the blueheaded wrasse, Thalassoma bifasciatum (Pisces: Labridae)[END_REF]. By removing large individuals via harvest, individuals change sex at an unnaturally small size. The consequences are many-fold. First the production of one type of gametes, for example, sperm in protogynous hermaphrodites, such as humphead wrasse and bumphead parrotfish, is lowered, since individuals assume the male role at a smaller size than natural, resulting in reduced fertilization success. Second, such smaller terminal phase males are less able to guard harems of females because they cannot fend off other males from taking over the harems [START_REF] Warner | Social control of sex change in the blueheaded wrasse, Thalassoma bifasciatum (Pisces: Labridae)[END_REF]. Likewise, larger females produce exponentially more, and often bigger, eggs that develop up to three times faster and survive starvation for more than twice as long as larvae produced by smaller females (Berkeley et al. 2004, Birkeland and[START_REF] Birkeland | The importance in fishery management of leaving the big ones[END_REF]. Thus, beyond the mere negative impacts of overfishing large individuals, in the case of sequential hermaphrodites, the removal of large individuals is in fact a selective removal of a key life history stage of the species that results in far reaching consequences (Armsworth 2001, Birkeland and[START_REF] Birkeland | The importance in fishery management of leaving the big ones[END_REF]. Importantly, while we discuss here only such impacts as a modification of fundamental life history traits, there are additional consequences at the level of the entire ecosystem, such as the overall health of coral reefs [START_REF] Mumby | Fishing, trophic cascades, and the process of grazing on coral reefs[END_REF][START_REF] Hughes | No-take areas, herbivory and coral reef resilience[END_REF], which directly impacts coral reef resilience [START_REF] Adam | Herbivory, connectivity, and ecosystem resilience: response of a coral reef to a large-Scale perturbation[END_REF].

Many commercial fishermen have traditionally targeted spawning aggregations. A number of fish families targeted by fishermen in the Indo-Pacific display spawning aggregations, groupers (Serranidae) and snappers (Lutjanidae) being the most studied, although other families, for example surgeonfishes (Acanthuridae) and rabbitfishes (Siganidae) also exhibit group spawning, but are less targeted [START_REF] Sadovy | Reef fish spawning aggregations: biology, research and management[END_REF]. Spawning aggregations are remarkable because they involve species that are usually found singly or in small groups, and are therefore not easy to catch in great numbers. During specific times of the year (usually related to moon phases), large numbers of individuals (thousands, sometimes tens of thousands) migrate to a single locale to spawn [START_REF] Meyer | Seasonal and diel movements of giant trevally Caranx ignobilis at remote Hawaiian atolls: implications for the design of marine protected areas[END_REF][START_REF] Erisman | Spatio-temporal dynamics of a fish spawning aggregation and its fishery in the Gulf of California[END_REF]. These locales remain the same over the years, allowing fishermen to predict and target the time and place of spawning. As mentioned above, fishing spawning aggregations is not a new activity; however, the advent of localization devices such as GPS has allowed the precise pinpointing of spawning grounds, something that traditionally prevented the extirpation of localized populations, in areas where spawning aggregations are not protected by artisanal fishermen [START_REF] Jokiel | Marine resource management in the Hawaiian Archipelago: the traditional Hawaiian system in relation to the Western approach[END_REF]. Removing large numbers of mature individuals at such a crucial period of the life cycle has obvious devastating effects. For example, the Caribbean Nassau grouper, Epinephelus striatus, which used to form aggregations of tens of thousands of individuals, are now considered endangered and most spawning aggregations have either disappeared entirely or have greatly diminished in numbers to the point where hundreds to a few thousand individuals may be seen [START_REF] Sala | Rapid decline of Nassau grouper spawning aggregations in Belize: fishery management and conservation needs[END_REF].

Case Studies for Applied Management and Conservation Integrating a Genetic Approach

Stock Delineation in the Context of Restoration: a Case Study from the Philippines.-Sea urchin fisheries are among the most lucrative shell fishery in some parts of the world, with Tripneustes gratilla being the most commercially important shell fishery in the tropical Indo-Pacific. The high market demand for export and unregulated harvesting of this species led to the collapse of the fishery in Bolinao, Pangasinan, NW Luzon, Philippines. Population genetic studies were undertaken to guide efforts to rebuild depleted spawning populations through release of hatchery produced juveniles [START_REF] Macawaris | Community-based sea urchin (Tripneustes gratilla) grow-out culture as a resource management tool[END_REF][START_REF] Ogden | Fisheries forensics: the use of DNA tools for improving compliance, traceability and enforcement in the fishing industry[END_REF]. Allozyme analysis detected no significant genetic structuring in the region [START_REF] Malay | Population genetics of the sea urchin Tripneustes gratilla[END_REF]. The population genetic structure of this species was re-investigated after the recovery of the fishery using mitochondrial and microsatellite markers [START_REF] Casilagan | Spatial and Temporal aspects of genetic variations in sea urchin Tripneustes gratilla populations in Western Luzon, Philippines[END_REF]. Neither marker detected significant genetic differentiation among populations in the northwest Luzon, nor among populations in regions where there has been no restocking and grow-out culture of hatchery produced sea urchins. Results of genetic studies concur with larval dispersal modeling studies, which indicated that T. gratilla populations in northwest Luzon were highly connected by larval transport due to the reversing monsoons (Juinio-Meñez and Villanoy 1995). Given the lack of dispersal barriers among localities and the spatio-temporal variability in recruitment due to hydro-meteorological factors, establishment of a regional network of marine protected areas and growout culture are strategic in conserving T. gratilla in the region (Juinio-Meñez et al. 2008). The high genetic diversity of the species despite recent recovery from depletion indicates the resilience of this species when populations are reestablished and sustainably managed.

Genetic Tools That Identify Both Sides of the Dispersal-Retention

Coin in Australian Reef Fishes.-Management actions need to include both the dispersal and retention elements of connectivity to ensure replenishment and ecological persistence in demographic timeframes, while also securing that genetic diversity can be maintained. Genetic diversity is important for several reasons, including the avoidance of inbreeding depression, the maintenance of healthy immune systems, and the maintenance of raw material for selection to act on different populations during times of environmental change [START_REF] Bowen | Gaia's handmaidens: the Orlog model for conservation biology[END_REF]. Here, we highlight two case studies of tropical Australian marine fishes that illustrate the important difference in dispersal over ecological and evolutionary time scales and the need to understand both in making management recommendations. Each case study is relevant to either conservation (the former) or fisheries (the latter) management. Each study, funded by a relevant industry stakeholder, will be incorporated into management strategies.

Connectivity and retention of the endemic anemonefish, Amphiprion mccullochi, living at the edge of south east Australian coral reef systems, was examined using mtDNA partial control region (D-loop) sequences and seventeen microsatellite DNA loci [START_REF] Van Der Meer | Historic hybridisation and introgression between two iconic Australian anemonefish and contemporary patterns of population connectivity[END_REF]. Extensive sharing of mtDNA haplotypes among sampling sites separated by up to 160 km indicates a lack of significant dispersal barriers over evolutionary time scales. In contrast, contemporary migration estimated using a Bayesian assignment technique indicated high levels of self-replenishment, with 68% to 84% of recruitment coming from within the same population [START_REF] Van Der Meer | Historic hybridisation and introgression between two iconic Australian anemonefish and contemporary patterns of population connectivity[END_REF]. The low proportion of dispersal among sites (1%-16%) indicates that populations are predominantly maintained by self-replenishment and should be managed separately, despite the lack of obvious barriers to dispersal through time seen in the mtDNA haplotype data. By understanding the interplay of self-recruitment and dispersal at various temporal and spatial scales, managers can seek to maintain self-seeding populations and high genetic diversity in this low abundance endemic species with a small geographic range and ecological specialization that make it more vulnerable to extinction.

In a counter example, the tropical Indo-West Pacific shorefish, Eleutheronema tetradactylus, from northern Australia was examined using partial mtDNA COI sequences and five microsatellite loci [START_REF] Horne | Limited ecological population connectivity suggests low demands on self-recruitment in a tropical inshore marine fish (Eleutheronema tetradactylum: Polynemidae)[END_REF]. Both microsatellite and mtDNA data indicated strong genetic structure existed among sites throughout northern Australia, suggesting that local populations are largely independent and insular, regardless of time scales. This inference was further substantiated for demographic timescales by non-genetic measures of connectivity in the same post-juvenile individuals, including: stable isotope profiles from sagittal otoliths [START_REF] Newman | Stock structure of blue threadfin Eleutheronema tetradactylum across northern Australia as inferred from stable isotopes in sagittal otolith carbonate[END_REF], parasite composition [START_REF] Moore | Stock structure of blue threadfin Eleutheronema tetradactylum across northern Australia, as indicated by parasites[END_REF], and life history characteristics [START_REF] Ballagh | Stock structure of the blue threadfin (Eleutheronema tetradactylum) across northern Australia derived from life-history characteristics[END_REF]. Together, these studies formed the basis for recommendations to establish regional management within each jurisdiction, cognizant of the resident nature of the populations.

Marine Protected Areas and Connectivity in South Africa.-South Africa is the meeting place of the Indian and Atlantic ocean and is bounded by two great ocean currents, the cold Benguela and the warmer Agulhas Current; with three distinct biogeographic regions in the area (Griffiths et al. 2010a). MPAs have been used globally to limit the impact of human activities on the marine environment, especially in the context of recreational and commercial exploitation [START_REF] Roberts | Effects of marine reserves on adjacent fisheries[END_REF][START_REF] Halpern | Marine reserves have rapid and lasting effects[END_REF][START_REF] Fernandes | Establishing representative no-take areas in the Great Barrier Reef: largescale Implementation of theory on marine protected areas[END_REF]. However, to be effective, it is recognized that networks of MPAs must be connected, either via adult or juvenile migration [START_REF] Botsford | Connectivity and resilience of coral reef metapopulations in marine protected areas: matching empirical efforts to predictive needs[END_REF]. Measuring connectivity using biological data is extremely difficult, but can involve tagging of migratory species or the use of otolith chemistry, trace-elemental fingerprinting, or trans-generational marking (see [START_REF] Levin | Review of methods for labelling and tracking marine invertebrate larvae[END_REF], 2006[START_REF] Thorrold | Transgenerational marking of embryonic otoliths in marine fishes using barium stable isotopes[END_REF][START_REF] Cowen | Larval dispersal and marine population connectivity[END_REF][START_REF] Carson | Evaluating the importance of demographic connectivity in a marine metapopulation[END_REF].

In South Africa, around 21% of the coastline is formally protected by MPAs, although only 9% are protected as "no-take" zones, in which all forms of exploitation are prohibited. Whether this network is effectively connected, especially across biogeographic boundaries remains unclear. Therefore a preliminary phylogeographic approach that compared multiple species, with different life-histories, across the same geographic range was used to determine connectivity between South African MPAs. Congruent genetic breaks across multiple species arise regardless of life history characteristics when there is a shared break or barrier to gene flow (Avise 1992, Kuo and[START_REF] Kuo | Phylogeographic breaks in low dispersal species: the emergence of concordance across gene trees[END_REF]. Therefore, a study that included twenty species (fishes, crustaceans, corals, echinoderms and molluscs) with different life history characteristics (broadcast spawning, brooding and live-bearing; see von der Heyden 2009) compared their population genetic patterns across this region. Most species analyzed showed at least one clear phylogeographic break across the range sampled, yet not all species showed the same breaks [START_REF] Teske | A review of marine phylogeography in southern Africa[END_REF]. A similar finding comes from a comparison of 27 species in Hawaii, where strong congruent patterns of genetic differentiation emerged, but no single species in the comparative analysis showed the emergent multispecies pattern [START_REF] Toonen | Defining boundaries for applying ecosystem-based management: a multispecies case study of marine connectivity across the Hawaiian Archipelago[END_REF]. Interestingly, dispersal estimates (calculated using isolation-by-distance approaches) of <10 km for brooding and live-bearing species are common (Wright et al. unpubl data).

How do genetics shed light on connectivity of MPAs in the region? First, the presence of at least four phylogeographic breaks that are shared between species suggests shared evolutionary processes, as has been found consistently in Indo-Pacific biogeographic studies as well (e.g., Lessios et al. 1996[START_REF] Gaither | Genetic evaluation of marine biogeographical barriers: perspectives from two widespread Indo-Pacific snappers (Lutjanus kasmira and Lutjanus fulvus)[END_REF], 2011). It therefore seems prudent to establish MPAs within each section of the coast between these phylogeographic breaks. Second, isolation-by-distance (IBD) analyses and population structuring indicate that successful dispersal might be far more limited than suggested by previous studies [START_REF] Brouwer | Adult movement and larval dispersal of Argyrozona argyrozona (Pisces : Sparidae) from a temperate marine protected area[END_REF][START_REF] Roberts | Currents along the Tsitsikamma coast, South Africa, and potential transport of squid paralarvae and ichthyoplankton[END_REF]. This strongly suggests that the current MPA network is ineffective at protecting genetic diversity (and adaptive potential). Therefore, evaluating genetic findings in combination with other parameters for establishing MPAs, such as abundance and diversity data, oceanographical and current patterns, and socio-economic data (see also [START_REF] Beger | Evolving coral reef conservation with genetic information[END_REF] will go some way in creating a more effective network in the region. Unfortunately, current scientific, funding, and political will are lacking in engaging with a genetic approach to management in the region. However, better science will inform stronger management-if genetics are increasingly used in marine conservation and management internationally, then South Africa will be placed in a strong position to build on existing genetic data.

Incorporating Genetics into Management Policy: Conservation in Hawaii.-Hawaii provides some of the better examples of genetic studies being incorporated directly into resource management policy. For example, population genetic studies of the endemic Hawaiian limpet, known locally as 'opihi (Cellana spp.), revealed that there was no detectable population differentiation within islands, but differentiation among islands [START_REF] Bird | Contrasting phylogeography in three endemic Hawaiian limpets (Cellana spp.) with similar life histories[END_REF]). Together with community involvement to survey and quantify population density differences among islands in the Hawaiian archipelago, this finding corroborated native Hawaiian practices to manage the stocks locally and has led to the introduction of several bills to change fisheries regulations for this species. Likewise, Christie et al. (2010a) tracked individual larval yellow tang, Zebrasoma flavescens, that were transported out of fisheries replenishment areas into adjacent regions open to collection along the Kona coast of the Big Island of Hawaii, documenting fisheries spillover from marine protected areas in Hawaii for the first time. This work revealed not only that spawning of abundant stocks in the replenishment area had benefits in the adjacent open areas, but also that long-lived pelagic larvae of these fishes recruit along the same coastline on which they were produced (Christie et al. 2010a). The recruitment of locally produced juveniles along the same coast was surprising to many, and has played a prominent role in the on-going debate about local management and the marine ornamental trade in Hawaii. Finally, a multispecies survey of population structure across the Hawaiian Archipelago revealed that the majority of 27 coral reef species surveyed showed genetic breaks between each of the adjacent Main Hawaiian Islands, despite the fact that they are geographically less separated than the more isolated Northwestern Hawaiian Islands (Toonen et al. 2011). Together, these studies have reformed the way that resource managers view populations of marine species in the Hawaiian Archipelago, resulting in a move away from archipelago-wide management strategies and a push for increased local management of marine resources. Efforts to communicate these findings widely are changing the type of policy being introduced by lawmakers as a result.

Increasing Public Ownership through Genetic Uniqueness in Fiji.-A population genetic survey revealed that Fijian regional color morphs of five putatively widespread fishes from the Indo-West Pacific (Amphiprion melanopus, Chrysiptera talboti, Pomacentrus moluccensis, Cirrhilabrus punctatus, and Labroides dimidiatus) were genetically differentiated from other locations with the typical color morph [START_REF] Drew | Endemism and regional color and genetic differences in five putatively cosmopolitan reef fishes[END_REF]. This information was something that excited local villagers and inspired a sense of nationalistic pride in the unique resources of the region. A public awareness and education campaign was launched, aimed at communicating that the people of Fiji are the stewards of their resources and that they are responsible for the health and survival of the world's entire population of these endemic fishes. The result has been an emerging realization of responsibility along with nationalistic pride, leading to the local phrase "Na ika kai viti," the "Fijian Fish" that has produced a widespread desire for conservation of these natural resources (J Drew, Columbia University, pers comm). With this sense of ownership and realized responsibility, environmental campaigns aimed at reducing anthropogenic impacts leading to coral reef decline, such as a campaign to control marine debris from the Fijian capital of Suva on the surrounding reefs, have become more prominent and well received.

Stock Structure and Cryptic Speciation in New Caledonian Mackerel (Scomberomorus spp.).-Improving knowledge of New Caledonian marine resources to increase sustainability is the global objective of the multidisciplinary program called ZoNéCo (Programme d'évaluation des resources marines de la zone économique de Nouvelle-Calédonie). Designed and launched >15 yrs ago by the Governments of France and New Caledonia and its three Provinces, together with locally-based research institutions, ZoNéCo acts as an interface linking science and policy, with the aim to support the implementation of research projects and to facilitate the uptake of research results by policy makers, the private sector, and civil society. Since its creation, the results produced and disseminated by the ZoNéCo program have led to a tremendous improvement in the knowledge of the marine ecosystems of New Caledonia.

Spanish mackerel, Scomberomorus commerson, is one of the most important commercially exploited and valuable fishes in New Caledonian waters and became the target of a major research program aimed at: (1) gathering knowledge on the ecology and exploitation of the species in the Indo-Pacific and New Caledonia, (2) monitoring biological parameters from populations throughout New Caledonia, and (3) determining management units within the archipelago. With this perspective, the genetic composition of S. commerson was investigated using mtDNA sequences (control-region and cytochrome-b gene fragments), and microsatellite allele frequencies at various spatial scales, from the local (Bélep in the North Province of New Caledonia) to the larger Indo-Pacific.

At the broadest scale, high levels of genetic divergence (>11%) were observed using mitochondrial sequences between Indo-Malay-Papua archipelago (IMPA) and both the southwestern Pacific, and the Persian Gulf and Oman Sea, suggesting the occurrence of two sister-species [START_REF] Fauvelot | Patterns of genetic isolation in a widely distributed pelagic fish, the narrow-barred Spanish mackerel (Scomberomorus commerson)[END_REF]. Further phylogeographic partitioning was evident between the western IMPA and the regions sampled east and south of it, i.e., northern Australia, West Papua, and the Coral Sea. Strong allele-frequency differences were found between local populations in the southwestern Pacific, both at the mtDNA (Φ st = 0.306-0.586) and microsatellite loci (West Papua vs New Caledonia: F ST = 0.202-0.311), indicating geographic isolation of S. commerson populations at this spatial scale [START_REF] Fauvelot | Patterns of genetic isolation in a widely distributed pelagic fish, the narrow-barred Spanish mackerel (Scomberomorus commerson)[END_REF].

Within the New Caledonia lagoon, nuclear data analyses revealed genetically distinct populations (F ST = 0.008-0.015; P < 0.01), one grouping samples from the East coast of New Caledonia and northern Bélep Islands, and another grouping samples from the west and south coasts of the main island (including Poya, Nouméa and Mont-Dore samples). The investigation of the temporal genetic composition of S. commerson captured in Bélep at the beginning of the year and at the end of the year were also genetically different (F ST = 0.008; P < 0.01). Although this genetic differentiation is low, it is significant and suggests that the Bélep Islands may host distinct reproductive stocks. The molecular data, which provides strong evidence for genetic structuring across a wide variety of spatial scales, is currently being discussed with policy makers to manage distinct stocks, and thus ensure long-term sustainable exploitation of this important resource.

Genetic Structure of Philippine Populations of Siganus fuscescens:

Implications for Fisheries Management.-Rabbitfishes (Family Siganidae) are a commercially valuable fishery and aquaculture species in the Philippines. Of the estimated 15 rabbitfish species which occur across the Philippine archipelago, the mottled spinefoot, Siganus fuscescens, is considered the most exploited, with evidence of overfishing [START_REF] Jumawan-Nanual | Population structure and reproductive biology of Siganus fuscescens Houttuyn 1782 (Perciformes, Siganidae) in Pujada Bay, Southeastern Mindanao, Philippines[END_REF]Metillo 2008, Soliman et al. 2009). Widely-distributed in reef flats and seagrass meadows, the species is harvested throughout most of its life cycle: presettlement juveniles are collected as they arrive in dense schools to settle on the reef flat, and juvenile and adult stages are fished year round [START_REF] Campos | Yield estimates, catch effort and fishery potential of the reef flat in Cape Bolinao, Philippines[END_REF]). Relatively much is known about the biology and ecology of the mottled spinefoot (e.g., [START_REF] Alcala | Ecological notes on rabbitfishes (Family Siganidae) and certain economically important marine animals in Southeastern Negros and environs, Philippines[END_REF][START_REF] De La Paz | Abundance and seasonality of siganid fishes (Teleostei, Perciformes) in Cape Bolinao, Pangasinan, Philippines[END_REF][START_REF] Woodland | Revision of the fish family Siganidae with descriptions of two new species and comments on distribution and biology[END_REF]), although some confusion still exists in distinguishing the species from its morphologically similar sibling species, Siganus canaliculatus. Several studies of mottled spinefoot populations in the Philippines have focused on population structure, dynamics, and exploitation status toward rationalizing and proposing management schemes for catch regulation [START_REF] Soliman | Assessment of the fishery of siganid juveniles caught by bagnet in Lagonoy Gulf, Southeastern Luzon, Philippines[END_REF] or establishment of marine reserves (Bellefleur 1997, Jumawan-Nanual and[START_REF] Jumawan-Nanual | Population structure and reproductive biology of Siganus fuscescens Houttuyn 1782 (Perciformes, Siganidae) in Pujada Bay, Southeastern Mindanao, Philippines[END_REF].

Recent genetic studies of Philippine populations of S. fuscescens have provided additional information which, coupled with earlier studies can be used to further inform management efforts. For example, phylogenetic studies have revealed the presence of cryptic diversity within the Philippines (Ravago-Gotanco and Juinio-Meñez 2010). While it remains unclear whether ecological partitions exist between the two distinct lineages, which occur in sympatry, these genetically distinct evolutionary units may be considered as potential management units (e.g., [START_REF] Bickford | Cryptic species as a window on diversity and conservation[END_REF][START_REF] Rocha | Phylogeography and the conservation of coral reef fishes[END_REF]). Likewise, significant population genetic structure along the eastern Philippine seaboard [START_REF] Magsino | The influence of contrasting life history traits and oceanic processes on genetic structuring of rabbitfish populations Siganus argenteus and Siganus fuscescens along the eastern Philippine coasts[END_REF], and among three major marine basin regions, i.e., South China Sea, northeast Philippine Sea, and southeast Philippine Sea-Sulu-Celebes Seas, indicate these should be considered as distinct regional management units (Ravago-Gotanco and Juinio-Meñez 2010). Across smaller spatial scales, analyses of mtDNA sequences and microsatellite data detect significant genetic structure across the western Luzon coast, relevant in particular for S. fuscescens populations in Bolinao, a significant fishery characterized by high levels of exploitation over many decades [START_REF] Campos | Yield estimates, catch effort and fishery potential of the reef flat in Cape Bolinao, Philippines[END_REF]. Assignment-based analyses of multilocus genotypes indicate asymmetric patterns of gene flow; populations in the Lingayen Gulf are self-recruiting, and serve as the source population for reefs located farther south along the western Luzon coast (Ravago-Gotanco 2010). Moreover, genetic estimates of gene flow are consistent with larval disperal estimates drawn from numerical modelling [START_REF] Altemerano | Influence of the Western Luzon Shelf Current on larval dispersal in Lingayen Gulf based on numerical Lagrangian dispersal experiments[END_REF]. While early maturity, high fecundity, short generation time, and the relative abundance of the species throughout the extensive seagrass habitats in the Bolinao-Anda reef system confer some degree of resilience, genetic studies suggest that local management and protection of Bolinao populations of S. fuscescens would benefit the entire western Luzon coast.

Summary

The present study highlights some of the ways in which molecular tools could be used to address conservation and management issues among marine taxa in the Indo-Pacific using examples of how such approaches have already been successfully applied. Molecular techniques and analyses allow us to examine not only the patterns of marine biodiversity within the region, but also test hypotheses regarding the processes that have shaped these patterns. Molecular tools provide invaluable insights to biological and evolutionary processes, but have been relatively slow to be incorporated into management. Failure to integrate genetic information risks losing the processes which maintain and generate biodiversity and potentially the populations and species of the future (Bowen andRoman 2005, Rocha et al. 2007). Most of the papers reviewed above have been based on mtDNA, but instead of single species, have often relied on multi-species approaches to look for congruent genetic breaks and patterns. Such an approach becomes even stronger when it includes species with different life history characteristics (see for example von der Heyden 2009, [START_REF] Toonen | Defining boundaries for applying ecosystem-based management: a multispecies case study of marine connectivity across the Hawaiian Archipelago[END_REF]. Multispecies approaches give far better resolution when elucidating the processes and patterns of marine biodiversity and can support integrative management and conservation goals. A major gap yet to be addressed is an integrated spatial planning approach which combines biological, oceanographic and socio-economic data with genetic data and evaluates spatial planning with and without taking evolutionary processes into account (see [START_REF] Beger | Evolving coral reef conservation with genetic information[END_REF]). An innovative and integrative approach is required to prioritize regions for conservation that can also incorporate uncertainty and take future change into account. highly variable and allow parent-off-spring relationships to be identified. Can be used to measure connectivity at contemporary timescales for better management than otherwise when using evolutionary tools. Need large numbers of adult samples and/ or recruits. Normally at small spatial scales, at single sampling time, so will not detect stochastic changes in space and time. Costs can be high for marker development.

Assignment/exclusion tests Same tools as for parentage analyses, but can be done on adult populations without requiring recruits. Is feasible with smaller adult samples than parentage, at greater spatial scales and at several times.

May underestimate connectivity and selfrecruitment among subpopulations with high gene flow.

Footnote: Definitions of terms-Connectivity, measures the amount of exchange between sub-populations; introns are non-coding parts of the nuclear genome; F st , is a population genetic measure of genetic differentiation between a subpopulation and the total population that it belongs to; ESU, Evolutionary Significant Unit, provides a rational basis for identifying genetically dicrete taxa that warrant priority treatment by conservation managers; MU, Management Unit, identifies the fundamental Unit for short-term management of populations with divergent allelle frequencies; Phylogeography, a bridge between macro-and micro evolution, which identifies coinciding geographic and genetic partitions across a species range; Parentage analysis, Genetic analysis that uses allelles present in offspring to assign them to their parent(s) in a sampled population. This most often uses microsatellite markers, but can also be performed on AFLP (Amplified Fragment Length Polymorphism) and SNP data; Assignment and exclusion tests: Genetic assignment methods permit estimates of contemporary dispersal by drawing inferences about where individuals were (or were not = exclusion) born based on their observed genotype likelihoods.

Although most population genetic approaches resolve some structure, there are numerous examples where genetic approaches are unable to resolve population level differentiation (Table 1). In such cases, it is often unclear whether the lack of resolution is a result of power, methodological limitations, or a true lack of any population differentiation [START_REF] Ryman | Power for detecting genetic divergence: differences between statistical methods and marker loci[END_REF][START_REF] Lowe | What can genetics tell us about population connectivity?[END_REF][START_REF] Marko | The complex analytical landscape of gene flow inference[END_REF]; Table 2). The emerging field of high-throughput next-generation sequencing shows extreme promise in facilitating broader phylogeographic studies (e.g., [START_REF] Puritz | Next-generation phylogeography: a targeted approach for multilocus sequencing of non-model organisms[END_REF], the evolution of mating systems (e.g., [START_REF] Hart | Next-generation studies of mating system evolution[END_REF], adaptation (e.g., [START_REF] Stapley | Adaptation genomics: the next generation[END_REF], and has been shown to resolve very recent divergence between marine and freshwater fish populations following an initial colonization event <150 yrs ago [START_REF] Czesny | Genetic divergence between freshwater and marine morphs of alewife (Alosa pseudoharengus): A next-generation sequencing analysis[END_REF]). In the Indo-Pacific region, next-generation sequencing has had a number of interesting applications and an exciting future; this is reviewed in more detail by [START_REF] Willette | So, you want to use next-generation sequencing in marine systems? insight from the Pan-Pacific Advanced Studies Institute[END_REF].

Finally, scientists need to communicate their work clearly to the general public as well as to the rest of the scientific and management community [START_REF] Olsen | Don't be such a scientist: talking substance in an age of style[END_REF][START_REF] Wilcox | It's time to e-Volve: taking responsibility for science communication in a digital age[END_REF]. Well-communicated science can influence conservation, policy, and ethical behavior, and bring excitement into school learning. Having a well-informed public could ultimately support future research when decisions are made on political leadership, who in turn decide to allocate funding to research, management, and conservation. We each have a novel and exciting story to tell about the beautiful marine environment we work in, which is probably one of the best ways of increasing global ownership and helping to conserve our amazing oceans.

Figure 1 .

 1 Figure1. Example of management units derived from overlaying genetic breaks occurring in three or more species (bold line). Management units in dashed lines are formed by breaks in fewer than two species, or where sampling was either incomplete, or lacking, or not covered by the literature reviewed here. Studies which formed part of this map are listed in Online Table1.

Table

  .

Table 1 .

 1 Example of molecular studies and species in the Indo-Pacific which resolve either population genetic structure or low/no structure. Studies such as these give critical insight into population connectivity over large geographical distances.

	Genetic structure

Table 2 .

 2 Summary of genetic tools used for management applications, together with a brief on the strengths and weaknesses of each. This is presented in two sections-evolutionary and ecological approaches. The footnote defines the terms used here and relevant references associated with the definitions given.Capacity to identify genetic breaks across many taxa for spatial management of ecosystems and/or target spp. As above and barriers are porous so vary over evolutionary time (e.g., sea level change creates or removes landbridges).

	Management applications

Acknowledgments

The authors would like to sincerely thank the National Evolutionary Synthesis Centre (NESCent) for hosting the Indo-Pacific Catalysis meeting, which brought together a diverse group of scientists working on the Indo-Pacific. Special thanks go to E Crandall and C Riginos for coordinating the meeting. This material is based upon work supported by the National Science Foundation through the National Evolutionary Synthesis Center (NESCent) under grant number NSF #EF-0905606. Thank you also to J Toms who helped with referencing and to J Gorospe for collating the species and authorities list.