The capture and culture of postlarval fish and invertebrates for the marine ornamental trade

Transcription

1 The capture and culture of postlarval fish and invertebrates for the marine ornamental trade JOHANN D. BELL 1, ERIC CLUA 2, CATHY A. HAIR 3, RENE GALZIN 4, PETER J. DOHERTY 5 1. Secretariat of the Pacific Community, B.P. D5, Noumea Cedex, New Caledonia 2. Coral Reef Initiatives for the Pacific, c/- Secretariat of the Pacific Community, B.P. D5, Noumea Cedex, New Caledonia 3. James Cook University, C/- Northern Fisheries Centre, PO Box 5396, Cairns, Queensland 4870, Australia 4. Ecole Pratique des Hautes Etudes ESA CNRS 8046, Universite de Perpignan, Perpignan Cedex, France 5. Australian Institute of Marine Science, PMB 3, Townsville, Queensland 4810, Australia Abstract Governments, non-government organizations and other stakeholders are striving to develop practices, policies and vehicles to make the tropical marine ornamental trade sustainable. Small-scale fisheries based on postlarval capture and culture (PCC) promise to contribute to this goal by (1) removing the risk of damaging corals (inherent in harvesting adults of target species established on reefs) by collecting postlarvae with light traps, nets and purpose-built temporary shelters as they settle from the plankton to the substrate; and (2) translating the high mortality of postlarvae at settlement into high rates of survival in culture. Possible concerns about overfishing of postlarvae, harvesting 1

2 the juveniles after they have run the gauntlet of predation at settlement, and the large proportion of bycatch, can be eliminated or greatly alleviated by restricting the size and quantity of fishing gear, designing it to retain bycatch alive, and releasing bycatch at times and places that minimize predation. However, special caution is needed when PCC is used at small, isolated islands with self-replenishing populations. Although PCC is environmentally friendly, its contribution to the ornamental trade is expected to be limited. Large variation in the abundance and species composition of settling postlarvae among years, the logistics and costs of operating labour-intensive operations in remote locations, and competition with responsible enterprises harvesting wild adults or producing ornamentals in hatcheries, are expected to constrain the viability and market share of dedicated PCC enterprises. PCC is expected to have the greatest uptake by parttime artisanal fishers in developing countries with infrastructure for exporting marine ornamentals. Such fishers are more immune to temporal variation in the supply of postlarvae - they can engage in PCC when valuable postlarvae are abundant and switch to other sources of income when they are scarce. Livelihood opportunities for smallholders could be enhanced through promotion of the environmental benefits of PCC among hobbyists maintaining marine ornamentals. Keywords coral reefs, sustainable small-scale fisheries, aquaculture, management, settlement of postlarvae 2

3 INTRODUCTION The trade in ornamental tropical marine fish and invertebrates has attracted much attention in recent years. At issue are the effects on coral reefs of collecting aquarium specimens, the livelihoods of the small-scale fishers in developing countries who supply the ornamental animals, and the interests of the millions of hobbyists who maintain tropical marine aquaria (Wood, 2001; Sadovy and Vincent, 2002: Green, 2003; Warbitz et al., 2003; McCollum, 2007; Reksodihardjo-Lilley and Lilley, 2007). The concerns arise not only because some of the species that contribute the 27 million fish, 16 million corals and 10 million other invertebrates traded each year may be over-exploited (Sadovy and Vincent, 2002), but also because destructive fishing methods are sometimes used to collect them (Green, 2003; Warbitz et al., 2003). The most notorious of these methods involves the illegal use of sodium cyanide to stun fish (Pet-Soede and Erdmann, 1998, Rubec et al., 2001), a practice that is also widespread in the live reef fish trade (LRFT) for food (Barber and Pratt, 1997a; Halim, 2002). Cyanide fishing has at least three undesirable consequences for coral reefs. First, a high proportion of fish die during collection, or shortly after purchase by hobbyists (Rubec, 1986; Hanawa et al., 1998; Rubec et al., 2001), with the result that more individuals are removed to meet demand than would be the case if more benign collection methods were used. Second, the physical structure of reefs is damaged when stunned fish retreat into holes and coral is broken apart to retrieve them (Halim, 2002). Third, there is collateral poisoning and death of non-target species of fish, corals and other invertebrates (Jones and Steven, 1997; McManus et al. 1997; Mous et al. 2000). However, high rates of mortality of marine ornamentals along the supply chain, and damage to corals, are not restricted to cyanide fishing; they can result from other 3

4 careless collection, husbandry and shipping practices (Rubec et al., 2001; Wood, 2001; Schmitt and Kunzman, 2005; Reksodihardjo-Lilley and Lilley, 2007). There are also serious concerns about the effects of sodium cyanide on the health of fishers and their families, and the tragedies of diving accidents and fatalities among poorly equipped and trained collectors (Jacques, 1997; Johannes and Djohani, 1997; Halim, 2002). Several non-governmental organizations (e.g., the International Marinelife Alliance, and the Haribon Foundation) and local government agencies have made earnest efforts to address the problems of destructive and unsafe fishing practices in the marine ornamental industry. In particular, they have: provided training for fishing communities in the use of barrier nets to collect fish without damaging corals (Anon. 1998; Rubec et al., 2001); identified the policy reforms and community-based management arrangements needed to establish incentives for cyanide-free aquarium and LRFT fisheries (Barber and Pratt, 1997b; Dalabajan, 2005); promoted higher prices for net-caught aquarium fish; and introduced testing to certify that specimens have not been collected using cyanide (Anon. 1998; Rubec et al., 2001, 2003). Regrettably, these efforts have only been partially successful. Reluctance on the part of many exporters and importers to pay higher prices for net-caught fish; the incentives of greater catches obtained through the use of cyanide, or simply the sheer difficulties in catching enough fish without using cyanide; and corrupt business practices continue to lock many fishers into this form of destructive fishing (Anon., 1998; Rubec et al., 2001; Lowe, 2002). The emerging framework for sustainable supply of marine ornamentals led by the International Marinelife Alliance and other NGOs has been strengthened by the creation of the Marine Aquarium Council (MAC) ( The aims of MAC are to establish standards for quality products and practices, certify compliance with these standards, 4

5 label the results of certification for quality assurance, and create consumer demand and confidence for certified and labeled organisms, practices and industry participants (Anon., 1999; Bunting et al., 2003). The MAC Core Standards for ornamentals collected from the wild address (1) ecosystem and fishery management, (2) collection, fishing and holding, and (3) handling, husbandry and transport. Together, they create a chain of custody from the reef to the home aquarium designed to assure consumers that they are purchasing high-quality ornamentals collected in a sustainable way. Ultimately, the success of certification systems like MAC depend on verifying that all parts of the process are implemented correctly (Shuman et al., 2004) and promoting the benefits to hobbyists (Alencastro et al., 2005; McCollum, 2007). Unless consumers are willing to pay more for eco-labeled ornamentals, the other participants further up the chain of custody will find it difficult to meet the costs of certification. Although widespread adoption of the MAC Core Standards for animals collected from the wild will undoubtedly help put the marine ornamental trade on a much more sustainable footing, it is not the only potential driver. Hobbyists are vitally concerned about the ease with which they can maintain fish and invertebrates in aquaria. Accordingly, they are prepared to pay a premium for cultured animals acclimated to life in captivity (Alencastro et al., 2005; McCollum, 2007). Thus, increasing the proportion of cultured ornamentals should help achieve the twin goals of reducing impacts on coral reefs and meeting the needs of consumers. It also has potential to create livelihoods (Job, 2005). Until recently, ornamentals cultured from hatchery-reared larvae accounted for only ~1% of the global trade (Wood, 2001). Such species were largely limited to (1) five or six species of giant clams (Heslinga et al., 1990, Lucas, 1994; Bell et al., 1997, Foyle et al., 5

6 1997; Hart et al., 1998, 1999), (2) a few species of shrimp (Warbnitz et al., 2003), and (3) fish with demersal eggs or mouth/pouch brooding habits and relatively large larvae, such as anemone fish (Pomacentridae), blennies (Blennidae), dottybacks (Pseudochromidae), gobies (Gobidae) and seahorses (Syngnathidae) (Ogawa and Brown, 2001; Olivetto et al., 2003; Warbnitz et al., 2003). However, advances in hatchery rearing, based on the use of copepods and SS-type rotifers as first food for the developing larvae, has paved the way for a wider range of fish species to be propagated, including the much sought-after angelfish (Pomacanthidae) (Cato and Brown, 2003; Job, 2005; Sim Sih Yang, pers. comm.). The willingness of consumers to pay 20-25% more for cultured marine ornamental fish because they have higher rates of survival in home aquaria (Alencastro et al., 2005; Job, 2005) is expected to provide incentives for increased hatchery production at a variety of scales. The collection and rearing of wild postlarvae, prior to and during settlement to coral reefs from the plankton, has also been proposed as a viable and sustainable method of supplying cultured ornamental fish and invertebrates (Dufour 2002; Hair et al., 2002a; Lecchini et al., 2006; Moana Initiative, 2007). Small-scale artisanal fisheries based on capture and culture of postlarvae promise to create livelihoods in developing countries, reduce the effects of the ornamental trade on coral reefs, and satisfy hobbyists. Here, we explain the possibilities for postlarval capture and culture (PCC); describe the main methods and benefits of PCC; assess the potential ecological impacts of collecting postlarvae; and outline the likely limitations and applications of PCC. Our review considers PCC only in the context of the ornamental trade. We do not generally discuss larger-scale aquaculture operations that collect and rear juvenile fish and 6

7 invertebrates at a range of sizes to supply markets for food, except to transfer lessons learned. POSSIBILITIES FOR POSTLARVAL CAPTURE AND CULTURE Three main factors make the capture and culture of postlarvae for the marine ornamental trade possible. First, the typical life cycle of tropical marine fish and invertebrates allows postlarvae to be collected relatively easily as they settle onto coral reefs. Second, a rapidly growing body of knowledge about the behaviour and ecology of postlarvae has revealed when and where postlarvae can be collected most efficiently. Third, the large differential in survival of settling postlarvae in the wild, and in culture, sets the stage for PCC to increase productivity without jeopardizing replenishment. These factors are explained in more detail below. Life cycle of coral reef fish and invertebrates Many of the animals living on coral reefs have a bipartite life cycle (Doherty, 1991; Leis, 1991). That is, the adults are usually associated with reefs, whereas the eggs and larvae are dispersed in the open sea (Figure 1). Typically, the gametes broadcast by highly fecund sedentary adults dwelling on reefs (Sale, 1980) spend weeks to months in the water column (Brothers and Thresher, 1985; Leis, 1991; Leis and McCormick, 2002). In the pelagic environment, the larvae develop through various stages until they metamorphose into postlarvae competent to settle onto reefs (Leis and McCormick, 2002; McCormick et al., 2002). Although <1% of larvae survive the pelagic phase to be potential colonists of reefs (Doherty, 1991; Leis, 1991), considerable numbers of robust postlarvae can return (Kami and Ikehara, 1976; Dufour et al., 1996; Letourneur et al., 1998; Dufour, 1999; Doherty, 2002; Doherty et al., 2004). When they do, they can be 7

8 caught alive from the water column near and within coral habitats using a variety of traps and nets (see below). Knowledge on behaviour and ecology of postlarvae An extensive literature on coral reef fish (reviewed in Sale, 2002) has helped identify the best times and places to catch juveniles settling from the plankton. Postlarvae of most demersal fish colonise reefs during nights around the new moon (Dufour and Galzin, 1993; Milicich and Doherty, 1994; Stobutzki and Bellwood, 1998; Leis and McCormick, 2002). Other studies (e.g., Reyns and Sponagule, 1999) have found the same timing for invertebrates. In all cases, settlement during the darkest hours is consistent with lower risk of detection by the wide range of visual predators associated with reefs. Synchronized settlement may also be a predator swamping strategy (Ims, 1990) facilitated by prey-switching predators (Tucker et al., 2008). Settlement of postlarvae around the new moon generally occurs throughout the year in the Caribbean (Robertson et al., 1993; Spongaule and Cowen, 1997; Wilson 2001) but is seasonal for fish and invertebrates in the Pacific (Talbot et al., 1978; Hair et al., 2002b, Hair and Doherty, 2003). Good locations for catching postlarvae are the outer slopes of reefs where they gather in preparation for settlement, in the passes or channels they use to enter lagoons (such as hoa in French Polynesia), and the reef crests and lagoons they traverse en route to the sheltered back-reef or fringing reef areas commonly inhabited by many species as juveniles. Knowing when and where to harvest postlarvae, however, is no guarantee that good catches will be made because there is great spatial and inter-annual variation in the availability of postlarvae. Typically, there are significant differences in the diversity and 8

9 abundance of settling postlarvae among reefs, and among years for any given reef (Doherty and Williams, 1988; Doherty, 1991: Doherty, 2002). Remarkably, for many species, the relative differences in abundance among reefs are generally maintained from year to year (Doherty and Fowler, 1994; Eggleston et al., 1998; Tolimieri et al., 1998, Booth et al., 2000; Doherty, 2002). Thus, a practical outcome for PCC from the research on spatial and temporal variation in abundance and diversity of postlarvae arriving on coral reefs has been identification of places that consistently have a greater supply of juveniles. Other such hotspots, at various scales, have been located by artisanal fishers, e.g., the areas now targeted by the communities catching and growing the puerulus larvae of spiny lobsters in Vietnam (Williams, 2004). The reasons for the distributions of settling postlarvae are complex, and the subject of continuing, vigorous research (for reviews see Cowan, 2002; Doherty, 2002; Leis and McCormick, 2002). In simple terms, the hotspots for the supply of postlarvae are probably created through the combined effects of (1) adaptations of fish and invertebrates to shed their eggs where larvae are likely to be entrained in currents that return them to suitable habitat (Cowan, 2002), (2) well-developed sensory systems and sophisticated behaviour of larvae and postlarvae, including strong swimming abilities, that help them use prevailing currents and features of reefs and their residents to return to natal areas (Stobutzki and Bellwood, 1997; Leis and Carson-Ewart, 1997; Swearer et al., 1999; Kingsford et al., 2002; Leis and McCormick, 2002; Myberg and Fuiman, 2002, Sympson et al., 2004; Leis, 2006; Montgommery et al., 2006; Lechini et al., 2007a; 2007b), and (3) differences in orientation and habitat quality among reefs (Doherty et al., 1996; Kingsford et al., 2002; Mellin et al., 2006). 9

10 Despite the accumulating knowledge about the factors determining the distribution of postlarvae, we are still a long way from predicting their abundance (Cowan, 2002). Part of the problem is that adaptations for larvae to return to natal reefs are often mediated or thwarted by the vagaries of currents, winds and storms (Cowen, 2002). Ultimately, predicting the abundance of postlarvae will involve modeling when the various factors favouring good supply (e.g., adequate food for the developing larvae, low abundances of predators, optimum climatic and oceanic conditions) are likely to coincide (Cowen, 2002). The cost of such modeling, and the necessary monitoring, is beyond the means of developing countries and the small-scale fishers expected to engage in PCC. Consequently, PCC will usually have to contend with considerable variation in catches of target species among years, even at the best sites. Differential rates of survival Although knowing where and when to find robust larvae is essential, the principal factor that has made PCC a possibility for the marine ornamental trade is the large differential in survival of settling postlarvae in the wild, and in culture. It has been known for some time that settling postlarvae experience high rates of predation ((Bailey and Houde, 1989; Doherty, 2002). Indeed, predation is thought to be so intense that postlarvae arriving on reefs have been said to face a wall of mouths (Hamner et al., 1988) and make a suicide drop onto the reef (Kaufman et al., 1992). This has been verified recently by a range of experimental studies (Planes and Lecaillon, 2001; Steele and Forrester, 2002; Webster, 2002; Doherty et al., 2004). Synthesis of mortality rates for 25 species involved in these and other studies by Almany and Webster (2006), and the study by Doherty et al. (2004), shows that a mean of 56.8% of juvenile coral reef fish are 10

11 consumed by predators within 1-2 days of settlement (Table 1). Substantial density independent and density dependent mortality of recently settled individuals then often occurs during the following weeks and months (Hixon and Webster, 2002). For example, total mortality of puerulus larvae during the first benthic year is 95-98% for several species of spiny lobster (Herrnkind and Butler, 1994; Butler et al., 1997; Phillips et al., 2003; Mills et al., 2004). The high mortality at settlement and during the first year of life is in stark contrast to the survival of the postlarvae if they are caught and reared in captivity. Wild-caught puerulus larvae of a variety of spiny lobster species have been reared for several weeks to 18 months with survival rates of 50-90% (Thomas et al., 2003; Williams, 2004; C. Hair, unpublished data). Durville et al. (2003) reared postlarvae of 10 species of coral reef fish for six months and obtained survival rates of 60-92%. Similarly, the Moana Initiative (2007) reports that 80% of target fish grown-out in PCC operations survive to the size demanded by the ornamental market, and Malpot et al. (in press) found that 98% of acanthurids survived after rearing for 60 days. Thus, provided fishing is limited so that it does not affect the potential for replenishment (see below), quarantining settling postlarvae and early benthic juveniles from predation through PCC represents a way to derive considerable additional production from ornamental species. In fact, the high rates of mortality at settlement, and the slow growth and longevity of many coral reef fish (Doherty and Fowler, 1994; Choat and Axe, 1996), allows a new paradigm to be considered for maximizing the sustainable harvest of some ornamental species, i.e., transferring fishing effort from adults to postlarvae (Bell et al. 1999; Hair et al., 2002; Hair and Doherty, 2003). In other words, for species suitable for PCC, harvesting and 11

12 rearing incoming settlers will be more sustainable than removing the same number of adults or older settled juveniles. MAIN METHODS FOR CAPTURING POSTLARVAE Although various techniques have been developed to harvest wild juvenile coral reef fish during their transition from the planktonic stage to the demersal stage at settlement (Choat et al., 1993), light traps and crest nets (Figure 2) are best suited for PCC. These two methods target postlarval fishes and are capable of retaining them alive in good condition. Light traps are submersible, passive devices that attract and concentrate zooplankton, including pre-settlement fishes (Doherty, 1987). Light traps can collect large numbers of postlarvae from relatively small volumes of water (Milicich, 1988), but are effective only for species that are photo-positive and relatively strong swimmers (Carleton and Doherty, 1997). Even so, they do not succeed in catching all the fish they attract, and there is some escapement of those fish that do enter traps (Meekan et al., 2000). To make light traps more suitable for PCC in developing countries, the original design has been modified to reduce costs (Watson et al., 2002). Some recent versions also provide the attracted fish with shelter (Lecaillon, 2004, Moana Initiative, 2007). The crest net was developed by Dufour and Galzin (1993) in French Polynesia and has now been used in several other countries, including Australia (Doherty and McIlwain 1996), Solomon Islands (Hair et al., 2002a) and La Reunion (Durville et al., 2003). Crest nets rarely exceed 3 m in mouth gape and can only be deployed at suitable locations, i.e., behind the surf zone on shallow reef tops with consistent unidirectional water flow. They capture postlarvae as they move at night from deeper waters outside reefs across the crest to settle in more sheltered backreef areas. Crest nets harvest a complementary range of ornamental species to light traps (Hair et 12

13 al., 2002b; Hair and Doherty, 2003), but catch a greater diversity and larger number of fish (Dufour et al., 1996). Provided crest nets are fitted with suitable chambers, a large proportion of the catch remains alive until the following morning. A variation on the crest net, the hoa net, is used to catch postlarvae in French Polynesia. This net takes it name from the location where it is installed; the shallow passes (hoa) that allow water to enter closed and semi-closed atoll lagoons (Lecaillon and Lourie, 2007). The mouth gape and collection chamber of the hoa net are similar to the crest net but it is fitted with large mesh wings to guide postlarvae into the mouth (Figure 3). A completely different device is used to collect postlarval spiny lobsters for the aquarium trade. It consists of a short section of a coconut log drilled with holes (Hair et al., 2007). These log traps are based on similar systems used in Vietnam to catch the settling puerulus larvae of Panulirus ornatus for aquaculture (Thuy and Ngoc, 2004), i.e., pieces of wood and dead coral drilled with holes. Log traps are anchored upright in shallow lagoonal areas behind reef crests (Figure 4). The settling puerulus larvae shelter in the holes overnight and can be removed the following day for grow-out in simple submerged containers (Hair et al., 2007). Although a range of other puerulus collectors have been used in Australia and New Zealand to monitor settlement for fisheries management purposes, and to obtain puerulus for aquaculture (Phillips and Booth, 1994; Phillips et al., 2001; Mills and Crear, 2004), they are generally designed for deeper water and are usually too expensive to be deployed by artisanal fishers in developing countries. 13

14 BENEFITS In addition to providing an opportunity to increase the productivity of target species by quarantining settling juveniles from predation, as outlined above, PCC has potential benefits for coral reefs, coastal fishing communities in developing countries, and the hobbyists who purchase ornamentals. The principal benefit for coral reefs is that the fish are collected from the water column away from the reef itself in the case of light traps, or in purpose built structures that can be installed away from areas of live coral in the case of crest and hoa nets, and log traps. Provided care is taken in setting and removing the traps and nets, PCC allows ornamentals to be collected without damaging corals, thus reducing one of the possible impacts of the marine ornamental trade on reefs. Coastal communities have the potential to benefit from PCC in several ways. First they can catch some specimens without having to use compressed air, thereby reducing the risk of diving accidents. Second, they can diversify the range of species they sell to the ornamental trade by supplying species that are otherwise difficult to find or collect at the desired size in the wild, e.g., spiny lobsters, or have distinctive juvenile colouration. Third, it allows small-scale operators to enter the market for cultured specimens, and benefit from the expected increase in demand for such animals, with modest investment compared to the cost of hatcheries. Another advantage is that the simple technology required for PCC enhances gender participation because some of the methods are more akin to reef gleaning than to fishing. The main advantage for hobbyists is that ornamentals produced by PCC are habituated to life in captivity because they are held in pens or tanks and fed for several 14

15 weeks. Therefore, they can be expected to have greater rates of survival in home aquaria relative to older specimens caught from the wild. Consumers will also have a wider choice of ornamentals because some species supplied by PCC cannot be collected effectively as adults in the wild, or produced in hatcheries. Like all forms of aquaculture, however, PCC operations may result in diseased animals without adequate husbandry. Training of villagers in maintenance of hygienic grow-out conditions is needed to minimize the risk of ornamentals from PCC contaminating the tanks of exporters, importers and hobbyists. POTENTIAL CONCERNS AND MEASURES TO MINIMIZE THEM Although the contributions that PCC can make to the marine ornamental trade hold much promise to be environmentally friendly, some concerns have been raised about the possible impacts of the methods. These concerns centre around the numbers and sizes of postlarvae collected, the bycatch, and the effects of removing postlarvae on the ecosystem. The nature of these concerns, their likelihoods and consequences, and the measures that can be applied to eliminate or minimize them are presented below and summarized in Table 2. The large numbers of postlarvae collected As more is learned about the great inherent spatial variation in the distribution of postlarvae settling on coral reefs (Doherty, 2002), and operators identify and focus on hotspots for settlement, there is concern that PCC could potentially result in overfishing under some circumstances. The concern arises because large collections of postlarvae (thousands to tens of thousands of individuals of a single species) have been made at one site during one night using crest nets and other devices (Dufour et al., 1996, Sadovy, 15

16 2001). Also, Lecaillon and Lourie (2007) report that hoa nets can catch millions of postlarvae from a variety of species. Thus, where postlarvae recruit at only a few limited, well known areas, at predictable times, intense fishing could remove a large proportion of them and jeopardize the replenishment of target species. Declines in abundance of other species collected as settling, or recently settled, juveniles for large-scale aquaculture, e.g., the fry of milkfish Chanos chanos (Ahmed et al., 2001), green grouper Epinephelus coioides (Johannes and Ogburn, 1999, see also Sadovy, 2001) and shrimp Penaeus monodon (Islam et al., 2004), certainly signal that overfishing of postlarvae may be possible. In some of these cases, however, the reduced catches cannot be attributed unequivocally to excessive harvesting; habitat degradation, pollution, and overfishing of adults are implicated as well. The consequences of removing the majority of a large cohort of postlarvae when it occurs are related to the fact that many coral reef fish are longlived (Choat and Axe, 1996) and strong year classes contribute a large proportion of spawning biomass (Doherty, 1991; Doherty and Fowler, 1994). Notwithstanding the density dependent and independent mortality associated with such strong year classes (Doherty, 2002; Hixon and Webster, 2002), fishing them down to the point where their contribution to the age structure of the population is equivalent to other cohorts can be expected to affect the potential for replenishment. As implied above, the likelihood that postlarvae can be overfished is largely restricted to small, isolated islands where populations of ornamental species are selfreplenishing (Cowen, 2002), and where it is easy to identify and target hotspots where the majority of postlarvae attempt to colonise. Such situations occur in Polynesia (see 16

17 Figure 5 for an example). In larger, more complex reef systems, the small size of light traps and crest nets relative to the large areas of coral reef means that the proportion of postlarvae captured by these methods is likely to be a tiny fraction of those recruiting. Indeed, the challenge for PCC in open reef systems will be to identify locations where sufficient postlarvae can be caught to make enterprises viable; see, for example, the low catch rates from light traps and crest nets in Australia and Solomon Islands (Doherty and McIlwain, 1996; Hair et al., 2002b). Even when hotspots for settlement of postlarvae are identified in large complex reef systems, the risk of overfishing a species is likely to be low because many fish and invertebrates form metapopulations in such ecosystems (Doherty, 2002; James et al. 2002; Kritzer and Sale, 2006). Metapopulations are typified by source populations, which not only replenish themselves and other components of the spawning stock, but also provide postlarvae for sink populations, where the output of adults to the spawning stock is insufficient to balance mortality (Figure 6). Metapopulations provide a measure of resilience against inadvertent overfishing of postlarvae - an overfished source area could be replenished from other sources, and an excessive harvest at a sink area has few consequences for the maintenance of effective spawning biomass. Although, PCC could potentially affect the replenishment of target species, there are several practical management measures that can be applied to eliminate or greatly reduce the risk of any such impacts. (1) For small, isolated islands, the number and types of nets and traps used, and the time and durations they are deployed, can be restricted to ensure that a substantial proportion of the postlarvae are not vulnerable to the gear. French Polynesia is proposing 17

18 such legislation to specify that hoa nets must not be deployed over more than two thirds of the width of the hoa and be limited to a maximum width of 20 m. In any event, largerscale PCC operators should be required to demonstrate that the proportion of the fish caught as postlarvae does not pose a risk to maintaining spawning biomass within sustainable bounds when combined with the catches of any existing fishery targeting adults. (2) Precautionary limits should also be placed on the numbers and sizes of fishing gear used for PCC on larger complex reefs. As information on metapopulation structure for ornamental species in such ecosystems accumulates, it should be used to locate PCC operations in areas that will have the least impact on the replenishment. (3) When strong year classes are encountered, and catches of target species exceed demand, the remainder of the individuals should be released into suitable habitat. (4) If concerns persist that harvests of postlarvae may be excessive, measures can be taken to make the effects of PCC biologically neutral at the very least. This approach stems from research underway in Australia to investigate the feasibility of collecting the puerulus larvae of spiny lobsters for aquaculture (Phillips et al., 2003; Gardner et al., 2006). For example, enterprises applying to catch puerulus of Jasus edwardsii are required to release 5% of the puerulus after they have been on-grown for one year to compensate for the percentage estimated to survive in the wild (Gardner et al., 2006). In addition, they are required to release another 20% of cultured one-year-old juveniles. The high rate of survival of puerulus in culture results in a win-win situation - the releases enhance the wild population supporting the capture fishery, and the large surplus of environmentally fit juveniles is available for aquaculture (Gardner et al., 2006). A 18

19 proviso is that the on-grown juveniles should be released at places and times that minimize predation - even when juveniles reach larger sizes, predation can pose a great obstacle to the success of stock enhancement initiatives (Bartley and Bell, 2008). Similar measures can be used when artificial shelters are deployed to collect postlarvae. Herrnkind et al. (1997, 1999) found that installation of structures with appropriate crevice sizes resulted in a 2- to 3-fold increase in settlement success of Panulirus argus in the Caribbean. Thus, removing settled individuals from a large subset of artificial shelters only would allow the remnant to enhance the wild population. Recommendations have been made to place such artificial structures for postlarvae where they do not rob natural habitats of potential recruits (Sadovy, 2001). However, use of physically attractive traps for settling postlarvae is analogous to the use of light traps and crest nets all methods collect animals that would have arrived on natural reefs but experience great predation there (Table 1). The risk of any impact of PCC on the replenishment of target species is also reduced by the nature of the ornamental trade. Unlike the large aquaculture industries based on collection of juvenile milkfish (Ahmed et al., 2001), groupers (Johannes and Ogburn, 1999; Sadovy, 2001, Ottolenghi et al., 2004) and spiny lobsters (Williams, 2004), the ornamental trade calls for relatively low numbers of a wide variety of species in each shipment (Green, 2003; Warbitz et al., 2003). Thus, demand for any given species from a PCC enterprise will be low, removing the incentive to catch large numbers of postlarvae. 19

20 Delaying the capture of settling fish The relatively rapid increase in the rate of survival of juvenile fish once they have run the gauntlet of predation at settlement (Doherty and Sale, 1985; Doherty, 2002, Almany and Webster, 2006) raises another potential concern delays in the capture of juveniles would make PCC more analogous to other forms of fishing (Sadovy and Pet, 1998, Mous et al., 2006). For example, Dufour (1999) has estimated that postponing capture of postlarvae from Day 5 to Day 50 after settlement increases the impact of fishing on the number of fish surviving to one year old by 3.5 times. The consequence of this is that the longer it takes to collect fish and invertebrates after they settle, the greater the effects of collection are on replenishment of the target species. The likelihood of PCC operations delaying the capture of settling postlarvae is not great, however. It is far easier to catch postlarvae while they are still in the water column or as they descend, than when they are already associated with corals and other benthic habitats. Indeed, light traps, crest nets, hoa nets and log traps have been designed to intercept postlarvae as they settle on reefs. These gears seldom catch larger individuals. Exceptions include some of the artificial shelters designed to catch groupers for the LRFT (Johannes and Ogburn, 1999; Mous et al., 2006). These structures attract settling postlarvae effectively, but older stages also move to them from nearby natural habitats. The potential impact of any PCC operations using artificial structures can be negated simply by removing postlarvae from the structures every few days and releasing any larger individuals. 20

21 Bycatch Another concern is that ornamental species are only a small proportion of the total catches made using most PCC methods, except for the log traps for puerulus larvae. For example, 94.7% of the fish collected by light traps and 97.7% of those collected in crest nets in Solomon Islands by Hair et al. (2002b) were either too small to be identified and cultured easily, or of no interest to the ornamental trade. Excessive bycatch has also been a feature of the larger fisheries based on the collection of juvenile shrimp (Islam et al., 2004), milkfish (Primavera, 2006) and groupers (Johannes and Ogburn, 1999; Sadovy, 1991; Ottolenghi et al., 2004; Mous et al., 2006) for aquaculture. In the case of the fisheries for shrimp and milkfish, billions of other organisms are collected and usually killed by the gear used to catch the target species, raising serious questions about the effects on the ecosystems involved (Primerva, 2006). While the scale of bycatch from PCC is orders of magnitude lower than for postlarval milkfish and shrimp fisheries, it must be handled responsibly if PCC is to be accepted and promoted as an environmentally friendly way of helping to supply the marine ornamental trade. The potential consequence of PCC for each bycatch species is the same as for the target ornamental species - replenishment could be jeopardised if harvesting is not limited, especially at small, isolated islands with self-replenishing populations (although, collectively the effects could be more profound because of the much greater number of species involved). Where the bycatch includes the juveniles of other economically valuable species, any such overfishing could affect the livelihoods of fishers who normally harvest these fish later in life for food or sale. Another potential consequence is 21

22 that removal of large numbers of postlarvae across a wide range of species may affect food webs on coral reefs. As described above for target species, the likelihood that any given bycatch species will be overharvested by PPC is low provided measures are taken to limit fishing at hotspots for recruitment on small, isolated islands. In reality, the likelihood is much lower because light traps and crest nets retain fish alive, which allows the bycatch to be released. Nevertheless, a number of measures need to be applied to PCC operations to ensure that mortality of bycatch is avoided or minimized. (1) The collection chambers for light traps, crest nets and hoa nets should be designed to retain fish and invertebrates in good condition. They should also be large and well-flushed enough to supply all fish with sufficient oxygen when strong year classes colonize reefs. This applies particularly to hoa nets, where simultaneous recruitment by many species has occasionally resulted in collection of millions of individuals, resulting in massive mortality due to overcrowding (Lecaillon and Lourie, 2007). Large catches of pomacentrids and clupeids in light traps can also cause high mortality (E. Malpot, pers. comm.). In the case of hoa nets, grids should be fitted across the gape of the net to prevent the entry of drifting algae, which can crush postlarvae in the retention chamber. Where jellyfish are common, light traps need to be modified to exclude them. (2) Light traps and nets deployed in the evening should be cleared at daybreak to minimize the time the catch is retained in the collection chamber. Systems should be developed to sort the target species on site so that the bycatch, including species that are too difficult to rear in captivity, can be returned to the water alive as quickly as possible. 22

23 However, where it is possible to retain the bycatch in a pen near the collection site, the fish should be released the following night to minimize predation. (3) If it is necessary to take part of the catch ashore for sorting, procedures should be developed to minimize stress and mortality of any remaining bycatch. Once sorted, it should then be released at the capture site that evening. (4) In the event that any of the bycatch dies, it should be distributed within the habitat where the fish would have settled so that it contributes to the food chain. Although this measure will not be totally effective, because some predators will only accept live food, the dead fish should provide food for a variety of scavenger species. Effects of removing target species The concern here is that even removing the small proportion of postlarvae of interest to the ornamental trade may deprive animal assemblages on coral reefs of some food. This potential impact needs to be weighed against the considerable environmental benefits of PCC, the interannual variation in the contribution of postlarvae to local food webs, and the likelihood that the assemblages of fish and invertebrates at the site are food limited. On balance, the impact on coral reef animal assemblages is unlikely to be greater than the widely accepted practice of fishing for adults. As Hair et al. (2002a) point out, the only difference is that removal of postlarval prey may have an effect on the condition, and therefore, egg production of stocks of piscivores, whereas removal of adults can spare prey species. Even so, any such effects are likely to be trivial due to the low total biomass of postlarvae collected. 23

24 LIMITATIONS Notwithstanding the development of methods for collecting postlarvae, and the potential for responsible PCC operations to make a sustainable contribution to the marine ornamental trade, the scope for PCC must be kept in perspective. A range of physical, environmental and economic constraints will limit establishment of viable operations, even at the scale of village-based artisanal fisheries. These limitations are outlined below. Selectivity and size of nets and traps The catches of postlarvae of interest to the marine aquarium trade from light traps and crest nets are often very limited due to the selectivity and small size of the gear. Consequently, even when harvests from both methods are combined, the numbers of valuable fish taken at any given location are well below the numbers of ornamental species exported for the region. For example, although ~130 ornamental fish species are exported from the Pacific to Canada alone (Baquero, 1999), only 57 and 27 species of value to the trade were caught in crest nets and light traps, respectively, in Solomon Islands (Hair et al., 2002b). Even then, only the subset of species of high-value were actually marketable due to the limited volume of airfreight from Solomon Islands (Hair and Doherty, 2003). Similarly, only 77 ornamental species were caught using crest nets in Moorea, French Polynesia (Lecchini et al., 2006). The catches from hoa nets can be an exception. When the net spans much of the width of a hoa, catches can be substantial (Lecaillon and Lourie, 2007) and good representation of the ornamental species present can be expected. However, most species in hoa are en route through these passes to lagoons; hoa nets do not catch species that 24

25 complete their life cycles within lagoons (Leis et al., 1998), or those distributed in deeper water as postlarvae (Leis, 1991) which settle on the outer reef slopes. The result of gear selectivity and gear size is that the proportion of the catch of potential value to the aquarium trade may be too low to make PCC profitable. For example, during experimental work to develop methods for PCC in Solomon Islands, Hair et al. (2002b) found that only 5.3% of fish caught in light traps were of value as ornamentals, of which ~99% were low-value pomacentrids. Similarly, 2.6% of fish and invertebrates in crest nets were of value, and 65% were pomacentrids. The fact that a species caught as a postlarvae is of medium- to high-value to the ornamental trade, does not always mean that this potential can be realized through PCC. Some potentially valuable postlarvae are not suitable for grow-out because they have specialized feeding habits and cannot be reared easily (Hair et al., 2002b). Spatial and temporal variation in supply of postlarvae Due to the relatively small size of the fishing gear, and the correspondingly low catch rates of valuable species, a key factor in determining the profitability of PCC will be locating hotspots that consistently receive relatively greater supplies of postlarvae. The work required to find such places is not too onerous for relatively small isolated islands where the self-replenishing populations have limited places to colonize back-reef areas, such as the closed and semi-closed atolls of the eastern Pacific. Nevertheless, sampling may be needed over several years to build up a reliable picture of the best areas to deploy the fishing gear due to the large-scale temporal variation of target species. For larger complex reef systems, like those in the western Pacific, considerably more time and expense will be involved in identifying locations where PCC is economically viable. 25

26 Such costs are likely to be beyond the resources of small-scale operators, who are more likely to deploy the gear on the nearest suitable, but perhaps non-productive, habitat. The inability to predict the supply of postlarvae of target species each year due to the significant temporal variation in abundance typical of many reef fish and invertebrates threatens the viability of larger-scale PCC operations. Lack of continuity in supply affects the ability of operators to fill orders and negotiate good prices. It also limits entry of small-scale fishers to those who can tolerate extended periods when valuable postlarvae are scarce. Market forces The share of the ornamental market that can be converted to PCC will be constrained by the inherent advantages of collecting wild specimens, and hatchery production. Table 3 summarises the features of species that are likely to be supplied from the wild capture fishery, PCC and hatchery-based aquaculture. Large juveniles and adults caught from the wild will continue to be the mainstay of a trade founded on a great diversity of animals (Wood, 2001; Warbitz et al., 2003), simply because this is the least expensive way to obtain the majority of species. The challenge is to implement the responsible chain of custody promoted by the Marine Aquarium Council to ensure that these animals are harvested in ways that do not damage coral reef ecosystems, or affect the replenishment of target species. Effort can be expected to be transferred from wild capture to PCC only in places where: i) overfishing of large juveniles/adults is deemed to be a risk; ii) postlarvae of target species can be obtained reasonably regularly and profitably by PCC; and iii) the species, or desired size of the species, is rare in the wild but the postlarvae are relatively common. 26

27 As profitable hatchery methods are developed for a wider variety of species, hatchery-reared animals can be expected to constrain the market for PCC products due to the inherent annual variation in supply of wild postlarvae. Both methods deliver acclimated ornamentals that attract premium prices but, where the costs of ornamentals produced by PCC and hatcheries are comparable, the trade would be expected to favour the hatchery-reared animals due to continuity of supply. The Marine Aquarium Council has now developed Mariculture and Aquaculture Management International Standards for cultured animals produced by hatcheries, and PCC ( As MAC standards are embraced more widely, and hatcheries become certified, PCC products will be at disadvantage if they do not also attain MAC certification. This will pose a problem for small-scale operators due to the expense involved, unless exporters cover the costs. In such situations, effective promotion by exporters of the environmental benefits of PCC is also likely to be needed to maintain market share. APPLICATIONS Taking these limitations into consideration, we expect PCC to contribute to the ornamental trade under two general circumstances. First, through full-time dedicated enterprises established at locations where high (but variable) supplies of postlarvae of medium-high value species, with good connections to export markets, result in total revenues well exceeding total costs. The profitability of such a hypothetical enterprise based in Mooera, French Polynesia, has been estimated by Lecchini et al. (2006). Other recent developments are described by the Moana Initiaitive (2007). 27

28 Second, through part-time activities of villagers who engage in PCC to diversify their income. Such enterprises will also need to have relatively easy access to export channels (Hair and Doherty, 2003), but would be expected to produce ornamentals intermittently, depending on the supply of postlarvae and other household activities. The scope for development of such small-scale enterprises is substantial in several developing countries, given the established infrastructure for the marine ornamental trade throughout much of the Indo-Pacific (Baquero, 1999; Wood, 2001; Warbitz et al., 2003). Hair and Doherty (2003) provide an example from Solomon Islands, where a small-scale operator using mainly crest nets and log traps would expect to be able to work fulltime on an attractive local income based mainly on the collection and rearing of ornamental shrimp and spiny lobsters. The manual on postlarval capture and grow-out by Hair et al. (2007) provides villagers with clear guidance on how to engage in the more simple forms of PCC. Other applications for PCC The great gains that can be made in the productivity of a species by catching it as postlarvae, and then rearing the juveniles in captivity to quarantine them from predation (Hair et al., 2002b), has been proposed as a way of restocking coral reefs (Moana Initiative, 2007). While there is little doubt that the capture, rearing and release involved can be done technically, it is dubious whether PCC has an important role to play as a tool for restocking. We say this because restocking is defined as release of cultured juveniles into wild population(s) to restore severely depleted spawning biomass to a level where it can once again provide regular, substantial yields (Bell et al., 2008). In fisheries where the number of adults is severely depleted it is improbable that postlarvae will be 28

29 abundant enough to catch, rear and release in numbers great enough to achieve the goals of augmenting spawning biomass substantially. Ultimately, restocking needs to demonstrate that it adds value to other forms of management (Bell et al., 2006). Otherwise, resources would be better placed into alternative forms of management, such as fully protecting the species and the habitats where juveniles recruit until the spawning biomass has reached the target level for reopening the fishery. Stock enhancement, which is defined as release of cultured juveniles into wild population(s) to augment the natural supply of juveniles and optimize harvests by overcoming recruitment limitation (Bell et al., 2008), has also been identified as a possible application for PCC. As outlined above, in the context of PCC, such releases are likely to be driven by concerns that harvests may affect replenishment. For stock enhancement to occur, the number of on-grown juveniles released would need to exceed the number required to make PCC biologically neutral. However, because effective stock enhancement often relies on releases of large numbers of cultured juveniles (Bell et al., 2005, 2006), the relatively small releases of on-grown juveniles possible through PCC operations are not expected to significantly augment the target self-replenishing populations of coral reef animals. Thus, whereas releases of a proportion of the individuals quarantined from predation may be a necessary part of responsible practice for PCC operations supplying the ornamental trade, the dedicated capture and culture of coral reef fish and invertebrates for stock enhancement is unlikely to be cost-effective. The scale of PCC is appropriate, however, for one potentially viable application for releasing on-grown postlarvae in the wild the enrichment of coral gardens at tourist destinations (Moana Initiative, 2007). Cultured juveniles can be released to fast- 29

30 track the colonization of fish, or establish the desired species composition of fish, on restored or constructed coral reefs at venues frequented by tourists. In such cases, PCC could be used to collect a wider range of species than those demanded by the ornamental trade. CONCLUSIONS The various methods for capturing postlarval fish and invertebrates alive as they settle to coral reefs has now been developed to the point where there is little doubt that they can supply a range of species for the ornamental trade in ways that: (1) remove the risk of damage to the physical structure of coral reefs, and (2) add substantial productivity to the target species by translating high mortalities at settlement into high rates of survival. Concerns that these benefits may be outweighed by possible negative effects on coral reef ecosystems can be largely allayed. The potential impacts of catching too many postlarvae, collecting them after they have passed through the survival bottleneck at settlement, and mortality of the large bycatch, can be eliminated or minimized due to the nature of PCC operations, and by responsible practice. In open, complex reef systems, the area fished by PCC gear is trivial compared to the area available for recruitment of postlarvae. Even when hotspots for recruitment are identified in such ecosystems, the metapopulation structure of most fish and invertebrates should often provide resilience against any inadvertent over-harvesting at one location. All PCC gear for ornamentals is designed to catch postlarvae as they migrate to and settle on reefs, removing the risk of capturing older individuals that have run the gauntlet of predation at settlement. Mortality 30

31 of bycatch can be avoided by well-designed PCC gear that retains settling juveniles alive, allowing them to be returned to the water at night when predation is lower. However, special care is needed for PCC operations on small, isolated islands with limited and well-known areas for recruitment of postlarvae. The risk of impacts is greater at such locations. Accordingly, limits must be placed on the sizes and numbers of fishing gear, and the times when they are used, to ensure that PCC does not jeopardize the potential for replenishment of target species, or result in excessive catches during periods of great postlarval abundance. The use of hoa nets needs particular scrutiny, responsible regulations and strict enforcement. We conclude that the main challenge for PCC is not environmental best practice but uptake. Considerable analysis will be needed to determine where PCC can be used to establish economically viable operations in competition with enterprises based on the responsible collection of older juveniles or adults from the wild, and juveniles produced in hatcheries. Such competition may limit larger-scale dedicated PPC operations to culturing those species that cannot be supplied by the other enterprises. The viability of such PCC operations will depend on identifying and obtaining access to hotspots for postlarvae, and contending with unpredictable supplies of medium- to high- value species due to temporal variation in recruitment of postlarvae, labour-intensive operations and the freight costs associated with remote locations. Premium prices for cultured and certified specimens promise to boost revenue, provided the benefits of PCC are promoted well, but the number of locations where dedicated larger-scale PCC operations will be profitable is expected to be limited. 31

32 On the other hand, PCC has potential to be adopted by a wide range of smallholders in countries where the infrastructure for export of ornamentals is well established. Smallholders will be more immune to variations in the supply of postlarvae they can engage in PCC when medium- to high- value postlarvae are relatively abundant and switch to a range of other income earning activities when the supply is poor. PCC can be embraced as a sustainable way of contributing to the marine ornamental trade, but potential investors, coastal communities, manager and NGOs should be realistic about the share of the market they can expect to capture, even with effective promotion of the products. ACKNOWLEDGEMENTS We thank Anne Gibert for assistance in obtaining the literature for this review, and Ian Hawes, David Lecchini, Emmanuel Malpot, Anne-Maree Schwarz, Antoine Teitelbaum and Beeing Yeeting for their helpful discussions about PCC or comments on the draft manuscript. REFERENCES Ahmed, M., G.A. Magnayon-Umali, R.A. Valmonte-Santos, J. Toledo, N. Lopez, and F. Torres, Jr. Bangus Fry Resource Assessment in the Philippines. ICLARM Technical Report No. 58. Manila: International Center for Living Aquatic Marine Resources Management (2001). Alencastro, L.A., R.L. Degner, and S. L. Larkin. Hobbyists preferences for marine ornamental fish: a discrete choice analysis of ecolabelling and selected product 32

33 attributes. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 15: (2005). Almany G.R., and M.S. Webster. The predation gauntlet: Early post-settlement mortality in reef fishes. Coral Reefs, 25: (2006). Anon. Haribon Netsman Program. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 4: 7-12 (1998). Anon. The Marine Aquarium Council, certifying quality and sustainability in the marine aquarium industry. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 5: (1999). Bailey, K.M., and E.D. Houde. Predation on eggs and larvae of marine fishes and the recruitment problem. Adv. Mar. Biol., 25: 1-83 (1989). Baquero, J. Marine Ornamentals Trade: Quality and Sustainability for the Pacific Region. Suva, Fiji: South Pacific Forum Secretariat (1999). Barber C.V., and V.R. Pratt. Sullied Seas: Strategies for Combating Cyanide Fishing in Southeast Asia and Beyond. World Resources Institute and International Marinelife Alliance (1997a). Barber, C.V., and V.R. Pratt. Policy reform and community-based programmes to combat cyanide fishing in Philippines. South Pacific Commission Live Reef Fish Info. Bull., 3: (1997b). Bartley, D.M., and J.D. Bell. Restocking, stock enhancement and sea ranching: Arenas of progress. Rev. Fish. Sci., 16: (2008). 33

34 Bell, J.D., L. Lane, M. Gervis, S. Soule, and H. Tafea Village-based farming of the giant clam, Tridacna gigas (L.) for the aquarium market: initial trials in the Solomon Islands. Aquacult. Res., 28: (1997). Bell, J.D., P. Doherty, and C. Hair. The capture and culture of postlarval coral reef fish: Potential for new artisanal fisheries. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 6: (1999). Bell, J.D., P.C. Rothlisberg, J.L. Munro, N.R. Loneragan, W.J. Nash, R.D. Ward, and N.L. Andrew. Restocking and stock enhancement of marine invertebrate fisheries. Adv. Mar. Biol., 49: (2005). Bell, J.D., D.M. Bartley, K. Lorenzen, and N.R. Loneragan. Restocking and stock enhancement of coastal fisheries: Potential, problems and progress. Fish. Res., 80: 1-8 (2006). Bell, J.D., K.M. Leber, H.L. Blankenship, N.R. Loneragan, and R. Masuda. A new era for restocking, stock enhancement and sea ranching of coastal fisheries resources. Rev. Fish. Sci., 16: 1-9 (2008). Booth D.J., M.J. Kingsford, P.J. Doherty, and G.A.Beretta. Recruitment of damselfishes in One Tree Island lagoon: Persistent interannual spatial patterns. Mar. Ecol. Prog. Ser., 202: (2000). Brothers, E. B., and R. E. Thresher. Pelagic duration, dispersal and the distribution of Indo-Pacific coral reef fishes. In: The Ecology of Deep and Shallow Coral Reefs, pp (M. L. Reaka, EWd.). NOAA symposium series on undersea research 3. Washington: United States Department of Commerce (1985). 34

35 Bunting, B.W., P. Holthus, and S. Spalding. The marine aquarium industry and marine conservation. In: Marine Ornamental Species: Collection, Culture and Conservation, pp (J.C. Cato and C. L. Brown, Eds.). Iowa: Iowa State Press/ Blackwell Publishing (2003). Butler, M. J., and W.F. Herrnkind. A test of recruitment limitation and the potential for artificial enhancement of spiny lobster (Panulirus argus) populations in Florida. Can. J. Fish. Aquat. Sci., 54: (1997). Butler, M. J., W. F. Herrnkind, and J. H. Hunt. Factors affecting the recruitment of juvenile Caribbean spiny lobsters dwelling in macroalgae. Bull. Mar. Sci., 61: 3 19 (1997). Carleton, J.H., and P.J. Doherty. The distribution and abundance of pelagic juvenile fish near Grub Reef, central Great Barrier Reef. Proc. 8 th Int. Coral Reef Symp., 2: (1997). Cato, J.C., and C. L. Brown (Eds.). Marine Ornamental Species: Collection, Culture and Conservation. Iowa: Iowa State Press/ Blackwell Publishing (2003). Choat, J.H., and L.M. Axe. Growth and longevity in acanthurid fishes: An analysis of otolith increments. Mar. Ecol. Prog. Ser., 134: (1996). Choat, J.H., P.J. Doherty, B.A. Kerrigan, and J.M. Leis. Sampling of larvae and pelagic stages of coral reef fishes: A comparison of towed nets, purse seine and light aggregation devices. Fish. Bull. U.S., 91: (1983). Cowen, R.K. Larval dispersal and retention and consequences for population connectivity. In: Coral Reef Fishes: Dynamics and Diversity in a Complex System, pp (P.F. Sale, Ed.). San Diego: Academic Press (2002). 35

36 Doherty, P.J. Light traps: Selective but useful devices for quantifying the distributions and abundances of larval fishes. Bull. Mar. Sci., 41: (1987). Doherty, P.J. Spatial and temporal patterns in recruitment. In: The Ecology of Fishes on Coral Reefs, pp (P.F. Sale, Ed.). San Diego: Academic Press (1991). Doherty, P.J. Variable replenishment and the dynamics of reef fish populations. In: Coral Reef Fishes: Dynamics and Diversity in a Complex System, pp (P.F. Sale, Ed.). San Diego: Academic Press (2002). Doherty, P.J., and P.F. Sale. Predation on juvenile coral reef fishes: an exclusion experiment. Coral Reefs, 4: (1985). Doherty, P.J., and D. McB. Williams. The replenishment of coral reef fish populations. Oceanogr. Mar. Biol., 26: (1988). Doherty, P.J., and A.J. Fowler. An empirical test of recruitment limitation in a coral reef fish. Science, 263: (1994). Doherty, P.J., and J. McIlwain. Monitoring larval fluxes through the surf zones of Australian coral reefs. Mar. Freshw. Res., 47: (1996). Doherty, P., M. Kingsford, D. Booth, and J. Carleton, J. Habitat selection before settlement by Pomacentrus coelestis. Mar. Freshw. Res., 47, (1996). Doherty, P.J., V. Dufour, R. Galzin, M.A. Hixon, M.G. Meekan, and S. Planes. High mortality during settlement is a population bottleneck for a tropical surgeonfish. Ecology, 85: (2004). Dufour, V. Population dynamics of coral reef fishes and the relative abundance of their early life history stage an example from French Polynesia. In: Proceedings of the Workshop on Aquaculture of Coral Reef Fishes and Sustainable Reef Fisheries, Kota 36

37 Kinabalu, Sabah, Malaysia, December 1996, pp (A.S. Cabanban and M. Phillips, Eds.). Kota Kinabalu, Sabah: Institute of Development Studies (1999). Dufour, V. Reef fish postlarvae collection and rearing programme for the aquarium market. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 10: (2002). Dufour, V., and R. Galzin. Colonization patterns of reef fish larvae to the lagoon at Moorea Island, French Polynesia. Mar. Ecol. Prog. Ser., 102: (1993). Dufour, V., Riclet, E., and A. Lo-Yat. Colonization of reef flat fishes at Moorea island, French Polynesia: Temporal and spatial variation of the larval flux. Mar. Freshw. Res., 47: (1996). Durville, P., P. Bosc, R. Galzin, and C. Conand. Aquacultural suitability of post-larval coral reef fish. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 11: (2003). Eggleston, D. B., R. N. Lipcius, L.S. Marshall, Jr., and S. G. Ratchford. Spatiotemporal variation in postlarval recruitment of the Caribbean spiny lobster in the central Bahamas: Lunar and seasonal periodicity, spatial coherence, and wind forcing. Mar. Ecol. Prog. Ser., 174: (1998). Foyle, T.P., J.D. Bell, M. Gervis, and I. Lane. Survival and growth of juvenile fluted giant clams, Tridacna squamosa, in large-scale village grow-out trials in the Solomon Islands. Aquaculture, 148: (1997). Gardner, C., S. Frusher, D. Mills, and M. Oliver. Simultaneous enhancement of rock lobster fisheries and provision of puerulus for aquaculture. Fish. Res., 80: Green, E. International trade in marine aquarium species: Using the global marine aquarium database. In: Marine Ornamental Species: Collection, Culture and 37

38 Conservation, pp (J.C. Cato and C. L. Brown, Eds.). Iowa: Iowa State Press/ Blackwell Publishing (2003). Hair, C., and P. Doherty. Progress report on the capture and culture of presettlement fish from Solomon Islands. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 11: (2003). Hair, C.A., J.D. Bell, and P. Doherty. The use of wild-caught juveniles in coastal aquaculture and its application to coral reef fishes. In: Responsible Marine Aquaculture, pp (R.R. Stickney and J.P. McVey, Eds.). New York: CAB International (2002a). Hair, C.A., P.J. Doherty, J.D. Bell, and M. Lam. Capture and culture of presettlement coral reef fishes in Solomon Islands. Proc. 9 th Int. Coral Reef Symp., (2002b). Hair, C., R. Warren, A. Tewaki, and R. Posalo. Postlarval Fish Capture and Grow-out. Canberra: Australian Centre for International Agricultural Research (2007). Halim, A. Adoption of cyanide fishing practice in Indonesia. Ocean Coast. Manage., 45: (2002). Hamner, W.M., M.S. Jones, J.H. Carleton, I.R. Hauri, and D.McB. Williams. Zooplankton, planktivorous fish, and water currents on a windward reef face: Great Barrier Reef, Australia. Bull. Mar. Sci., 42: (1988). Hanawa, M., L. Harris, M. Graham, A. Farrell, and L. Bendall-Young. Effects of cyanide exposure on Dascyllus aruanus, a tropical marine fish species: Lethality, anesthesia and physiological effects. Aqua. Sci. Conserv., 2: (1998). 38

39 Hart, A.M., J.D. Bell, and T.P. Foyle. Growth and survival of the giant clams Tridacna derasa, T. maxima and T. crocea at village farms. Aquaculture, 165 (3/4): (1998). Hart, A.M., J.D. Bell, I. Lane and T.P. Foyle. Improving culture techniques for villagebased farming of giant clams (Tridacnidae). Aquacult. Res., 30: (1999). Herrnkind, W. F., and M. J. Butler. Settlement of spiny lobster Panulirus argus (Latreille, 1804), in Florida: Pattern without predictability? Crustaceana, 67: (1994). Herrnkind, W. F., M.J. Butler, J.H. Hunt, and M. Childress. Role of physical refugia: Implications from a mass sponge die off in a lobster nursery in Florida. Mar. Freshw. Res., 48: (1997). Herrnkind, W. F., M.J. Butler, and J.H. Hunt. A case for shelter replacement in a disturbed spiny lobster nursery in Florida: Why basic research had to come first. Am. Fish. Soc. Symp., 22: (1999). Heslinga, G.A., T.C. Watson, and T. Isamu. Giant clam farming. Honolulu: Pacific Fisheries Development Foundation (NMFS/NOAA) (1990). Hixon, M.A., and M.S. Webster. Density dependence in reef fish populations. In: Coral Reef Fishes: Dynamics and Diversity in a Complex System, pp (P.F. Sale, Ed.). San Diego: Academic Press (2002). Houde, E.D. Comparative growth, mortality, and energetics of marine fish larvae: temperature and implied latitunidal effects. Fish. Bull. U.S., 87: (1989). Ims, R.A. On the adaptive value of reproductive synchrony as a predator-swamping strategy. Am. Nat., 136: (1990). 39

40 Islam, M.D., M.A. Wahab, and M. Tanaka. Seed supply for coastal brackishwater shrimp farming: Environmental impacts and sustainability. Mar. Poll. Bull., 48: 7-11 (2004). Jacques, M. Doomed fishermen. South Pacific Commission Live Reef Fish Info. Bull., 3: (1997). James, M.K., P.R. Armsworth, L.B. Mason, and L. Bode. The structure of reef fish metapopulations: modeling the larval dispersal and retention patterns. Proc. R. Soc. Lond. Ser. B, 269: (2002). Job, S. Integrating marine conservation and sustainable development: Communitybased aquaculture of marine aquarium fish. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 13: (2005). Johannes, R.E., and R. Djohani. Reducing the incidence of the bends in Indonesian fishing villages: Education may not be enough. South Pacific Commission Live Reef Fish Info. Bull., 3: 40 (1997). Johannes, R.E., and N.J. Ogburn. Collecting grouper seed for aquaculture in the Philippines. Secretariat of the Pacific Community Live Reef Fish Info. Bull.6: (1999). Jones, R.J., and A.L. Steven. Effects of cyanide on corals in relation to cyanide fishing on reefs. Mar. Freshw. Res., 48: (1997). Kami, H.T., and I.I. Ikehara. Notes on the annual juvenile siganid harvest in Guam. Micronesica, 12:

41 Kaufman, L., J. Ebersole, J. Beets, and C.C. McIvor. A key phase in the recruitment dynamics of coral reef fishes: Post-settlement transition. Environ. Biol. Fish., 34: (1992). Kingsford, M.J., J.M. Leis, A. Shanks, K.C. Lindeman, S.G. Morgan, and J. Pineda. Sensory environments, larval abilities and local self-recruitment. Bull. Mar. Sci., 70(1) SUPPL: (2002). Kritzer, J.P., and P.F. Sale. The metapopulation ecology of coral reef fishes. In: Marine Metapopulations, pp (J.P. Kritzer and P.F. Sale, Eds.). San Diego: Academic Press (2006). Lecaillon, G. The C.A.R.E. (collect by artificial reef eco-friendly) system as a method of producing farmed marine animals for the aquarium market: An alternative solution to collection in the wild. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 12: (2004). Lecaillon, G., and S. M. Lourie. Current status of marine postlarval collection: Existing tools, initial results, market opportunities and prospects. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 17: 3-10 (2007). Lecchini, D., S. Polti, Y. Nakamura, P. Mosconi, M. Tsuchiya, G. Remoissenet, and S. Planes. New perspectives on aquarium fish trade. Fish. Sci., 72: (2006). Lecchini, D., S. Planes, and R. Galzin. The influence of habitat characteristics and conspecifics on attraction and survival of coral reef fish juveniles. J. Exp. Mar. Biol. Ecol., 341 : (2007a). 41

42 Lecchini, D., C.W.O. Osenberg, J.S. Shima, C. St Mary, and R. Galzin. Ontogenetic changes in habitat selection during settlement in a coral reef fish: ecological determinants and sensory mechanisms. Coral Reefs, 26: (2007b). Leis, J.M. The pelagic stage of reef fishes: The larval biology of coral reef fishes In: The Ecology of Fishes on Coral Reefs, pp (P.F. Sale, Ed.). San Diego: Academic Press (1991). Leis, J.M. Are larvae of demersal fishes plankton or nekton? Adv. Mar. Biol., 51: (2006). Leis J.M., and B.M. Carson-Ewart. In situ swimming speeds of the late pelagic larvae of some Indo-Pacific coral-reef fishes. Mar. Ecol. Prog. Ser., 159: (1997). Leis J.M., and B.M. Carson-Ewart. Complex behaviour by coral-reef fish larvae in open water and near-reef pelagic environments. Environ. Biol. Fish., 53: (1998). Leis, J.M., and M. I. McCormick. The biology, behaviour and ecology of the pelagic, larval phase of coral reef fishes. In: Coral Reef Fishes: Dynamics and Diversity in a Complex System, pp (P.F. Sale, Ed.). San Diego: Academic Press (2002). Leis, J.M., T. Trinski, P.J. Doherty, and V. Dufour. Replenishment of fish populations in the enclosed lagoon of Taiaro Atoll (Tuamoto Archipelago, French Polynesia): Evidence from eggs and larvae. Coral Reefs, 17: 1-8 (1998). Letourneur, Y., P. Chabanet, L. Vigliola, and M. Harmelin-Vivien. Mass settlement and post-settlement mortality of Epinephelus merra (Pisces: Serranidae) on reunion coral reefs. J. Mar. Biol. Assoc. U. K., 78: (1998). Lipcius, R.N., D.B. Eggleston, S.J. Schreiber, R.D. Seitz, J. Shen, M. Sisson, W.T. Stockhausen, and H.V. Wang. Importance of metapopulation connectivity to restocking and restoration of marine species. Rev. Fish. Sci., 16: (2008). 42

43 Lowe, C. Who is to balme?: Logics of responsibility in the live reef food fish trade in Sulawesi, Indonesia. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 10: 7-16 (2002). Lucas, J.S. The biology, exploitation and mariculture of giant clams (Tridacnidae). Rev. Fish. Sci., 2: (1994). McCollum, B.A. Consumer perspectives on the web of causality within the marine aquarium trade. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 17: (2007). McCormick, M., L. Makey, and V. Dufour. Comparative study of metamorphosis in tropical reef fishes. Mar. Biol., 141: (2002). McManus J.W., R.B. Reyes, Jr., and C.L. Nanola, Jr. Effects of some destructive fishing methods on coral cover and potential rates of recovery. Environ. Manage., 21: (1997). Malpot, E., R. Galzin, and G. Remoissenet. Utilization des larves de poisons de recifs coralline : Synthese des travaux menes en Polynesie Francaise. Secretariat of the Pacific Community Live Reef Fish Info. Bull., (in press). Meekan, M.G., P.J. Doherty, and L. White Jr. Recapture experiments show the low sampling efficiency of light traps. Bull. Mar. Sci., 67: (2000). Milicich, M.J. The distribution and abundance of presettlement fish in the nearshore waters of Lizard Island. Proc. 6 th Int. Coral Reef Symp., 2: (1988). 43

44 Milicich, M.J., and P.J. Doherty. Larval supply of coral reef fish populations: Magnitude and synchrony of replenishment to Lizard island, Great Barrier Reef. Mar. Ecol. Prog. Ser., 110: (1994). Mills, D., and B. Crear. Developing a cost-effective puerulus collector for the southern rocl lobster (Jasus edwarsdii) aquaculture industry. Aquacult. Eng., 31: 1-15 (2004). Mills, D. J., C. Gardner, and S. Ibbott. Behaviour of on-grown juvenile spiny lobsters, Jasus edwardsii, after re-seeding to a coastal reef in Tasmania, Australia. In: Stock Enhancement and Sea Ranching: Developments, Pitfalls and Opportunities, pp , (K. M. Leber, S. Kitada, L. Blankenship and T. Svasand, Eds.). Oxford: Blackwell Publishing (2004). Mellin, C., J. Ferraris, R. Galzin, M. Kulbicki, and D. Ponton. Diversity of coral reef fish assemblages: Modelling of the species richness spectra from multi-scale environmental variable in the Tuamotu Archipelago (French Polynesia). Ecol. Model., 198: (2006). Moana Initiative. La PCC: Un outil pour la Conservation et la Valorisation de la Biodiversite. (www. moanainitiative.org) (2007). Montgomery, J.C., A. Jeff, S.D. Simpson, M.G. Meekan, and C. Tindle. Sound as an orientation clue for the pelagic larvae of reef fish and crustaceans. Adv. Mar. Biol., 51: (2006). Mous P.J., L. Pet-Soede, M. Erdmann, H.S.J. Cesar, Y. Sadovy, and J.S. Pet. Cyanide fishing on Indonesian coral reefs for the live food fish market what is the problem? Secretariat of the Pacific Community Live Reef Fish Info. Bull., 7: (2000). 44

45 Mous, P.J., Y. Sadovy, A. Halim, and J. S. Pet. Capture for culture: Artificial shelters for grouper collection in SE Asia. Fish and Fisheries, 7: (2006). Myberg, Jr., A.A., and L.A. Fuiman. The sensonry world of coral reef fishes. In: Coral Reef Fishes: Dynamics and Diversity in a Complex System, pp (P.F. Sale, Ed.). San Diego: Academic Press (2002). Ogawa, T., and C. Brown. Ornamental fish aquaculture and collection in Hawaii. Aqua. Sci. Conserv., 3: (2001). Oliveto, I., M. Cardinali, L. Barbaresi, F. Maradonna, and O. Carnevali. Coral reef fish breeding: The secrets of each species. Aquaculture, 224: (2003). Ottolenghi, F., C. Silvestri, P. Giordano, A. Lovatelli, and M.B. New. Capture-based Aquaculture: The Fattening of Eels, Groupers, Tunas and Yellowtail. Rome: Food and Agriculture Organization of the United Nations (2004). Pet-Soede, L., and M. Erdmann. An overview and comparison of destructive fishing practices in Indonesia. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 4: (1998). Phillips, B. F. and J. D. Booth. Design, use and effectiveness of collectors for catching the puerulus stage of spiny lobsters. Rev. Fish. Sci. 2: (1994). Phillips, B. F., R. Melville-Smith, Y. W. Cheng, and M. Rossbach. Testing collector designs for commercial harvesting of western rock lobster (Panulirus cygnus) puerulus. Mar. Freshw. Res., 52: (2001). Phillips, B.F., R. Melville-Smith, and Y.W. Chung. Estimating the effects of removing Panulirus cygnus pueruli on the fishery stock. Fish. Res., 65: (2003). Planes, S., and G. Lecaillon. Caging experiment to examine mortality during metamorphosis of coral reef fish larvae. Coral Reefs, 20: (2001). 45

46 Primavera, J.H. Overcoming the impacts of aquaculture on the coastal zone. Ocean Coast. Manag., 49: (2006). Reksodihardjo-Lilley, G., and R. Lilley. Towards a sustainable marine aquarium trade: An Indonesian perspective. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 17: (2007). Reyns, N., and S. Sponaugle. Patterns and processes of brachyuran crab settlement to Caribbean coral reefs. Mar. Ecol. Prog. Ser., 185: (1999). Robertson, D.R., U.M. Schober, and J.D. Brawn. Comparative variation in spawning output and juvenile recruitment of some Caribbean reef fishes. Mar. Ecol. Prog. Ser., 94: (1993). Rubec, P.J. The effects of sodium cyanide on coral reefs and marine fish in the Philippines. In: The First Asian Fisheries Forum, pp (J.L Mclean, L.B. Dizon and L.V. Hosillos, Eds.). Manila: Asian Fisheries Society (1986). Rubec, P.J., F. Cruz, V. Pratt, R. Oellers, B. McMullough, and F. Lallo. Cyanide-free netcaught fish for the marine aquarium trade. Aqua. Sci. Conserv., 3: (2001). Rubec, P.J., V.R. Pratt, B. McCullough, B. Manipula, J. Alban, T. Espero, and E.R. Supildo. Trends determined by cyanide testing on marine aquarium fish in the Philippines. In: Marine Ornamental Species: Collection, Culture and Conservation, pp (J.C. Cato and C. L. Brown, Eds.). Iowa: Iowa State Press/ Blackwell Publishing (2003). Sadovy, Y. Summary of regional survey of fry/fingerling supply for grouper mariculture in Southeast Asia. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 8: (2001). 46

47 Sadovy, Y., and J. Pet. Wild collection of juveniles for grouper mariculture: Just another capture fishery? Secretariat of the Pacific Community Live Reef Fish Info. Bull., 4: (1998). Sadovy, Y.J., and A.C.J. Vincent. Ecological issues and the trades in live reef fishes. In: Coral Reef Fishes: Dynamics and Diversity in a Complex System, pp (P.F. Sale, Ed.). San Diego: Academic Press (2002). Sale, P.F. The ecology of fishes on coral reefs. Oceanogr. Mar. Biol. Ann. Rev., 18: (1980). Sale, P.F. Introduction. In: The Ecology of Fishes on Coral Reefs, pp (P.F. Sale, Ed.). San Diego: Academic Press (1991). Sale, P.F. Coral Reef Fishes: Dynamics and Diversity in a Complex System. San Diego: Academic Press (2002). Schmitt, C., and A. Kunzman. Post-harvest mortality in the marine aquarium trade: A case study of an Indonesian export facility. Secretariat of the Pacific Community Live Reef Fish Info. Bull., 13: 3-12 (2005). Shuman, C.S., G. Hodgson, and R.F. Ambrose. Managing the marine aquarium trade: is eco-certification the answer? Environ. Conserv., 31: (2004). Simpson, S.D., M.G. Meekan, R.D. McCaugley, and A. Jeff. Attraction of settlementstage coral reef fishes to reef noise. Mar. Ecol. Prog. Ser., 276: (2006). Swearer, S.E., J.E. Caselle, D.W. Lea, and R.R. Warner. Larval retention and recruitment in an island population of a coral-reef fish. Nature, 402: (1999). Sponaugle, S., and R.K. Cowen. Early life history traits and recruitment patterns of Caribbean wrasses (Labridae). Ecol. Monogr., 67: (1997). Steele, M.A., and G.E. Forrester. Early postsettlement predation on three reef fishes: 47

48 Effects on spatial patterns of recruitment. Ecology, 83: (2002). Stobutzki, I.C., and D.R. Bellwood. Sustained swimming abilities of the late pelagic stages of coral reef fishes. Mar. Ecol. Prog. Ser., 149: (1997). Stobutzki, I.C., and D.R. Bellwood. Nocturnal orientation to reefs by late stage coral reef fishes. Coral Reefs, 17: (1998). Talbot, F.H., B.C. Russell, and G.R.V. Anderson. Coral reef fish communities: Unstable, high-diversity systems? Ecol. Monogr., 48: (1978). Thomas, C. W., C. G. Carter, and B. J. Crear. Feed availability and its relationship to survival, growth, dominance and the agonistic behaviour of the southern rock lobster, Jasus edwardsii in captivity. Aquaculture, 215: (2003). Thuy, N.T.B., and N.B. Ngoc. Current status and exploitation of wild lobsters in Vietnamese waters. In: Spiny Lobster Ecology and Exploitation in the South China Sea Region. Pp (K.C. Williams, Ed.). Canberra: Australian Centre for International Agricultural Research, Proceedings No. 120 (2004). Tolimieri, N., P.F. Sale, R.S. Nemeth, K.B. Gestring. Replenishment of populations of Caribbean reef fishes: are spatial patterns of recruitment consistent through time? J. Exp. Mar. Biol. Ecol., 230: (1998). Tucker, J.K., G.L. Paukstis, and F.J. Janzen. Does predator swamping promote synchronous emergence of turtle hatchlings among nests? Behav. Ecol. 19: (2008). Warbitz, C., M. Taylor, E. Green and T. Razak. From Ocean to Aquarium. Cambridge: UNEP- World Conservation Monitoring Centre (2003). 48

49 Watson, M., R. Power, S. Simpson, and J.L. Munro. Low cost light traps for coral reef fishery research and sustainable ornamental fisheries. Naga, the ICLARM Quarterly, 25(2): 4-7 (2002). Webster, M.S. Role of predators in the early post-settlement demography of coral-reef fishes. Oecologia, 131: (2002). Williams, K.C. Spiny Lobster Ecology and Exploitation in the South China Sea Region. Canberra: Australian Centre for International Agricultural Research, Proceedings No. 120 (2004). Wilson, D.T. Patterns of replenishment of coral-reef fishes in the nearshore waters of the San Blas Archipelago, Caribbean Panama. Mar. Biol., 139: (2001). Wood, E.M. Collection of Coral Reef Fish for Aquaria: Global Trade, Conservation Issues and Management Strategies. Herefordshire, UK: Marine Conservation Society (2001). 49

50 LIST OF FIGURE CAPTIONS Figure 1. The bipartite life cycle of coral reef fish and invertebrates, where the adults are associated with coral reef habitats and the pelagic eggs and larvae develop in the ocean. [See Sale (1991) for variations on this general theme]. Figure 2. a,b) Light traps used to catch photo-trophic postlarval coral reef fish, and (c,d) crest nets used to capture postlarval fish and invertebrates traversing reef crests to settle in sheltered back reef areas. Figure 3. The hoa net used to catch postlarval fish colonizing atoll lagoons in French Polynesia (Photo courtesy of Christophe Brie). Figure 4. a, b) Drilled coconut logs used to catch the puerulus larvae of spiny lobsters in Solomon Islands, and c) juvenile Panulirus versicolor reared for 3 weeks following collection from log traps (Photos courtesy of C. Hair). Figure 5. Ahunui Atoll in French Polynesia, an example of a small isolated island with a closed lagoon and limited places (hoa) where postlarvae can recruit (see arrows). (Photo courtesy of Serge Andréfouët). Figure 6. Source and sink populations within meta-populations (redrawn from Lipcius et al., 2008). 50

51 Table 1. Summary of the mortality of 25 fish species within 2 days of settling on coral reefs (after Almany and Webster, 2006). Family Species Mortlaity (%) Days Acanthuridae Acanthurus sp Acanthurus triostegus Naso unicornis* 61 <1 Zebrasoma scopas Acanthurus coeruleus 47 1 Balistidae Rhinecanthus aculeatus Chaetodontidae Chaetodon auriga 92 2 Chaetodon ephippium Chaetodon citrinellus Gobiidae Coryphopterus glaucofraenum 26 1 Coryphopterus nicholsi 92 1 Gnatholepis thompsoni 6 1 Pomacentridae Dischistodus perspicillatus 92 2 Hemiglyphidodon plagiometapon 38 2 Neopomacentrus azysron 50 2 Neopomacentrus cyanomos 44 2 Pomacentrus amboinensis 62 2 Pomacentrus nagasakiensis 85 2 Pomacentrus pavo 31 2 Stegastes albifasciatus Stegastes nigricans Chromis viridis Chrysiptera leucopoma Stegastes leucostictus 63 1 Stegastes partitus 69 1 Average = 56.8 * Data from Doherty et al. (2004) 51

52 Table 2. Summary of the potential concerns about the effects of postlarval capture and culture for the ornamental trade on coral reef ecosystems, together with their possible consequences and likelihoods, the measures that can be taken to eliminate or minimize them, and an assessment of the overall risk (see text for details). Potential Concern Consequence Likelihood Measures to address concern Risk* Too many Reduced potential for Possible for small, isolated islands Limit numbers and sizes of traps and nets, and times/durations of deployment Low postlarvae are replenishment of target Negligible for open, complex Release large cohorts surplus to demand collected species reefs Release a proportion of on-grown juveniles to achieve biological neutrality Delaying the capture of settling fish Mortality of the large bycatch Reduced potential for replenishment of target species. Reduced potential for replenishment of bycatch species Alters local foodwebs if bycatch is removed Low because most PCC gear targets settling postlarvae artificial shelter traps for fish are the exception Low because gear is designed to keep postlarvae alive Collect postlarvae from artificial shelter traps every few days and release any older individuals Ensure collection chambers are big enough to keep all fish alive if large collections are made Remove postlarvae from gear early in the morning, sort bycatch on site, retain alive and release at night Distribute dead bycatch in settlement habitat Low Low Removal of target postlarvae affects food webs Condition/egg production of predators is reduced Low because total catch of target postlarvae is usually small. Release fish from large cohorts surplus to demand Low *Assuming measures are applied responsibly 52

53 Table 3. Features of marine ornamental fish and vagile invertebrate species likely to determine the way they are supplied to the tropical marine aquarium trade. Note that this analysis doe not include invertebrates like corals, giant clams and echinoderms. Source of Supply Features of species Example families/genera Collection of wild adults/large juveniles Capture and culture of postlarvae Hatcheries Low-medium value, easy and inexpensive to collect High value, possible to collect easily High value, difficult to catch and rear as postlarvae, and to produce in hatcheries Medium-high value, easy to collect and rear as postlarvae but difficult to catch in the wild or produce in hatcheries At risk of overfishing as adults, but available and easy to rear as postlarvae Medium-high value, life cycle can be closed and hatchery production is profitable Pomacentrids Labridae Pomacanthidae Acanthuridae Paniluridae Stenopus Tetraodontidae Acanthuridae Balistidae Chaetodontidae Serranidae Amphiprion Gobiidae Pomacanthidae Pseudochromis 53

54 Figure 1 Adults Settlers Colonization Recruits Eggs Postlarvae Larvae Dispersion 54

55 Figure 2 (a) (credit Anthonin Hubert) (b) (credit Sandrine Polti) (c) (credit WorldFish Center) (d) (credit CRIOBE) 55

56 Figure 3 56

57 Figure 4 a) b) c) people science environment partners 57

58 Figure 5 58