CHAPTER 8

Shallow Water Reef-associated Finfish

Andrew Wright

I. INTRODUCTION

Throughout the western and South Pacific, reef-associated finfish form the basis of most subsistence fisheries. In many areas, this resource is also fished on a commercial scale. Although user conflicts resulting from the sharing of this resource are well known, studies detailing social and resource interaction issues that develop with increased fishing are limited. Compounded by a poor understanding of social factors affecting fishing in coastal communities throughout the Pacific is a generally poor but increasing knowledge of the population biology and fishery dynamics of target finfish resources. At least this is the case from the perspective of western science. Current knowledge of the biology and population dynamics of many fish species targeted by small-scale fisheries in the South Pacific region is poor relative to that available for the site-attached component of the reef fish community found in shallow water. This component, which is relatively easy to study, comprises mainly small species that are generally planktivorous or herbivorous and school, is currently only lightly exploited throughout the South Pacific region for the aquarium trade (see Pyle, this volume). Documented knowledge of the biology and population dynamics of food fish such as the scarids, lethrinids, lutjanids, carangids and the serranids, has improved significantly during the last decade. Contributing to the rapid in­ crease in knowledge of these fish is the fact that these families are generally circum-tropical in distribution. Thus information recorded throughout their range assists in providing a fundamental understanding of the biology of the same species in the South Pacific. In the short- to medium-term, fisheries managers in the South Pacific islands countries will continue to rely on such sources of information. At present, local research is generally not well pre­ pared, in terms of financial and manpower resources, to undertake comprehen­ sive studies of fish biology which could support fisheries development and management throughout the region. 204 Andrew Wright

However, among many South Pacific coastal communities, there exists a depth of understanding of marine resources and their environment that has been accumulated over centuries of observation and which is passed orally among generations. In few instances has this reservoir of knowledge and associated traditional practice been applied to contemporary fisheries development and management issues. Documentation of this information which commenced in the late 1970s in Palau (Johannes, 1977,1978a, 1981 a) has since been expanded throughout the region (e.g. Johannes, 1982a and b; Hviding, 1990; Sims, 1990, Zann, 1990, Polunin, 1990; Baines, 1990; Wright, 1990). The South Pacific presents some unique opportunities for innovative man­ agement of fisheries development. It is a region where western concepts and aspirations increasingly impinge upon traditional values and practices. It is a region not only where politics are playing an increasing role in resource use decisions but also where rural communities utilise the traditional knowledge base to extract maximum, but not necessarily sustainable, benefit from resource exploitation. This chapter reviews information that is available on the biology of tropical reef-associated finfish that are taken by fisheries operating in the islands of the western and South Pacific. In many instances it is complemented by chapters elsewhere in this volume on small schooling pelagic fishes, deepwater reef slope finfish resources and aquarium fishes. It commences with an overview of the zoogeography and taxonomy of the major family groups considered, outlining difficulties experienced by biologists in islands countries with identification of species. This is followed by discussion of habitats associated with reef systems in the western and central Pacific and factors affecting finfish diversity. The following section is concerned with the study of diets and trophic relationships of finfish in reef-associated environments. Methods for the study of reproduc­ tive biology, mortality and growth and the conclusions of research on these topics are then discussed. A brief description of fisheries operating in the South Pacific, including gears and catch composition, precedes a final section which introduces issues associated with attempts to manage multispecies fisheries for finfish among islands communities in the South Pacific. In most instances the information has been retrieved from published sources and in these situations, full citations are given. A comprehensive bibliography accompanies this review to facilitate the work of fisheries researchers in the region. Not all these references have been cited in the text; they are provided as a guide for further reading and a starting point for research. In addition, citations have largely been limited to those in the English literature. There is also a significant amount of information, albeit largely inaccessible to many researchers in Pacific islands countries, in the Chinese, French, Soviet and Japanese literature. Shallow Water Reef-associated Finfish 205

II. LIFE HISTORY AND POPULATION BIOLOGY

It is probable that there are between 6,000 and 7,000 species of fishes distributed through two classes, Osteichthyes and Chondrichthyes, throughout the Indo-Pacific faunal region. The 20 to 30 species of cartilagineous fishes, some of which are of interest to fisheries, are discussed elsewhere (Nichols, this volume). This chapter generalises across 2,000 species of ray-finned fishes (Actinopterygii) found in association with coral reefs, or approximately half the total shore fish fauna recorded from the Indo-Pacific region and one-sixth the total known to science. It is the 300 or more species representing 30 to 50 families taken by subsistence or commercial fisheries that are of primary interest here. Dalzell (this volume) discusses the biology and fisheries for small schooling pelagics associated with coral reefs, Moffit (this volume) discusses those species found in deeper water on reef slopes and Pyle (this volume) discusses the smaller species generally not targeted by food fisheries but which are commercially valuable in the international aquarium trade.

ZOOGEOGRAPHY

Although this review is primarily concerned with the island groups of Melanesia, Micronesia and Polynesia, the geographical distribution of many of the species of finfish considered in this chapter extends throughout the tropical Indo-Pacific. The largest diversity of tropical marine fishes is found in the southern Philippines and eastern Indonesia where 90 per cent of all shore fishes recorded from the Indo-Pacific are found (Myers, 1989). Within the western Pacific region, 165familiescomprisingmorethan2,500specieshavebeenrecorded. The diversity of the nearshore fish fauna decreases east and west of this region (Sale, 1980; Springer, 1982; Woodland, 1986). As noted by Myers (1989), although field material facilitating a comparison of fish diversity across the tropical Pacific is generally lacking, available data illustrates how dramatic the change in species diversity is (Fig. 1). There are a large number of fish species found in Indonesia and the Philippines which do not occur to the east; 151 reef species do not occur beyond Yap or Palau and 231 reef species do not occur east of the eastern Carolines (Myers, 1989). This is related to a number of biotic and abiotic factors including the apparent frequency of environmental disturbance (Sale, 1980; Springer, 1982; Rosenblatt and Waples, 1986), geological history, the diversity of habitats available in the mid-South Pacific with a decrease in the occurrence of high islands and an increasing prevalence of atoll-type environments eastwards. On the basis of life history strategies, Kulbicki (1992) analysed community struc- / Ryukyus 1 Hawaiian Islands >1,209 /.' 460

J) Philippines Marianas 872 \ Johnston Is. fc5? Belau-Yap 271 , "': Marshalls . 827 j y Papua New Gu e. Carolines

".. Marqu I Society IIs . /' ca- 35

/ " , ; -. ' P

Rapa 220

Figure 1. The southern and western Pacific region illustrating recent estimates for the num region and likely routes of colonisation (after Myers, 1989). Shallow Water Reef-associated Finfish 207 ture for a range of reef systems across the Pacific. Based on a division of life history characteristics ranging from small, short-lived, relatively highly fecund species that attain maturity early in their life cycle to large, late maturing, long- lived species, he described two major groups of reef fishes in the region: 1) a western and central Pacific group which comprised three sub-groups, the West Pacific, central Pacific and the Great Barrier Reef-New Caledonian regions and 2) an eastern and southern Pacific group sub-divided into Hawaiian, Polynesian and Southern regions. These groupings mirror well the limits of tectonic plates in the Pacific region. The number of species of scarids and siganids, for example, decreases across the central Pacific to the east. Whereas the greatest number of scarid species, approximately 30, is recorded in the Philippine and Indonesian Archipelagoes, and the adjacent western Pacific, the tropical eastern Pacific supports only six species, three of which are endemic (Choat and Randall, 1986). Similarly for siganids, thirteen species are reported from the Philippines, 10-12 from Mela­ nesia, 6 from Micronesia and only 2 from Western Samoa in the east (Woodland, 1983). Eachfaunal area, the Red Sea-Indian Ocean, Indonesia-Philippines and the eastern Pacific, contain a mixture of area-specific and wide ranging species. Although larval dispersal capabilities are not well understood (Montgomery, 1990), it has been suggested that many species have larval lives that are too short to enable them to survive extended periods of pelagic drift required to colonise distant reef environments (Springer, 1982). Thus, the high islands in the southeastern region of the South Pacific do not receive many larval colonisers from the species-diverse sources in the western Pacific. In a north-south direction, current movements are not conducive to the transport of fish larvae, with the result that only 1,200 species of fish have been recorded from the Ryukyus in southern Japan and approximately 900 from the southern Great Barrier Reef. Recommended further reading on the zoogeography of fishes in the western and South Pacific includes Mead (1970), Briggs (1974), Springer (1982), Myers (1989) and Montgomery (1990).

TAXONOMY

Despite the paramount importance of species identification to research concerning fish biology and the development of fisheries management and species conservation advice, one of the major problems facing fisheries workers in the South Pacific region concerns accurate identification offish. Although this is not a problem unique to the South Pacific (e.g. Beliwood, 1988a), it continues to result in wasted utilisation of scarce research funds throughout the region. Modern taxonomic keys are often difficult to obtain and publications that collate available information in an easily referable format are generally expen- 208 Andrew Wright

Table I. Useful literature to assist with the identification offish commonly harvested by South Pacific fisheries.

Family Reference

Acanthuridae Randall (1955a; 1955b; 1956; 1960a; 1987a), Shen and Lim (1973) Albulidae Shaklee and Tamaru (1981) Balistidae Matsuura (1979; 1980), Randall and Klausewitz (1973), Randall etal. (1978), Hutchins (1986) Belonidae Mees (1962), Parin (1967), Carangidae Suzuki (1962), Matsui (1967), Smith (1967-1970; 1972), Beny and Cohen (1972), Berry etal. (1983), Gushiken (1983), Williams etal. (1980), Williams and Venkataramani (1978) Congridae Kanazawa (1958), Smith (1962), Castle (1968) Coryphaena Palko etal. (1982) Cynoglossidae Menon (1977; 1979) Diodontidae Leis(1978) Exocoetidae Parin (1961a; 1961b; 1967a) Fistularidae Fritzche (1976) Haemulidae Lee (1985) Hemiramphidae Collette and Parin (1978), Parin el al. (1980) Holocentridae Greenfield (1974), Shimizu and Yamakawa (1979), Randal and Gueze (1981), Matsuura and Shimizu (1982), Randall etal. (1982), Allen and Cross (1983) Kyphosidae Tsutsumi and Kurata (1971), Fujita et al. (1984) Labridae Schultz(1960) Leiognathidae James (1975) Lethrinidae Smith (1959), Sato (1978) Lutjanidae Schultz (1953a), Anderson, Kami and Johnston (1977), Anderson, Talwar and Johnston (1977), Johnson (1980), Anderson (1981), Akazaki (1983), Allen (1985), Allen and Talbot (1985), Carpenter(1987), Kishimoto etal. (1987), Raj and Seeto (1983) Mugilidae Thompson (1954), Pillay (1962a; 1962b) Mullidae Lachner(1954; 1960), Shinohara (1967), Randall (1974), Yoshino and Sata (1981), Shen (1976) Muraenidae Rosenblatt and McCosker(1971), McCoskerand Rosenblatt (1975), Randall and McCosker(1975), Hatooka (1982) Nemipteridae Wongratana (1974; 1978), Rao and Rao (1981) Platycephalidae Matsubara and Ochiai (1955), Schultz (1966), Wongratana (1975), Shao and Chen(1987) Plectropomids Hoese etal. (1981), Randall and Hoese (1986) Sauridae Shinda and Yamada (1972) Scaridae Schultz (1958; 1969), Randall (1963), Randall and Bruce (1983), Bruce and Randall (1985), Choatand Randall (1986) Scomberomorus Collette and Russo (1978; 1980) Scombridae Collette and Russo (1983), Collette and Nauen (1983), Collette (1983a) Scorpaenidae Smith (1957a; 1957b) Serranidae Schultz (1953b), Randal] (1964; 1987b), Morgans (1965), Gosline (1966), Zama (1978), Randall and Ben-Tuvia (1983), Randall and Heemstra (1986;1991), Randall and Hoese (1986) Siganidae Lam (1974), Burhanuddin etal. (1975), Woodland (1983) Soleidae Shen and Lee (1981) Sphyraenidaede Sylva (1973), Au (1979) Teraponidae Vari (1978) Shallow Water Reef-associated Finfish 209 si ve. In the last few years a number of excellent publications covering all known genera have been produced which, although limited in geographic coverage, provide useful assistance for a great number of species distributed throughout the tropical Indo-Pacific (see Table I and II). Good examples of this include Fourmanoir and Laboute (1976), Grant (1978), Masuda et al. (1984), Gloerfelt- Tarp and Kailola (1984), Sainsbury et al. (1985), Myers (1989) and Randall et al. (1990). Although such publications generally do not lend themselves well to use in the field, they have resulted in a significant improvement in taxonomic reference material available to fisheries workers in the region. They improve substantially upon reference material available in earlier but significant publi­ cations on South Pacific fishes such as Fowler (1928; 1931; 1934; 1949), Munro (1967) and the Species Identification Guides produced by the Food and Agriculture Organisation (FAO) for the central western Pacific (Fischer and Whitehead, 1974). In addition to reference material which consolidates taxonomic information for all genera, new species, new information on species distribution and

Table II. Useful literature to assist with the identification of fish in the South Pacific region.

Region Reference

Fiji Fowler (1959) French Polynesia Randall (1964; 1973) Great Barrier Reef Burgess and Axelrod (1975b), Grant (1978), Russell (1983), Randall etal (1990) Guam Kami et al. (1968), Ikehara (1970), Kami (1971; 1975), Jones and Chase (1975), Myers and Shepard (1980), Amesbury aand Myers (1982) Johnston Island Gosline (1955), Randall etal. (1985) Kiribati Schultz (1943), Randall (1955c), Chave and Eckert (1974) Lord Howe Island Talbot and Whitley (1976) Marianas Myers (1988) Marshall Islands Randall (1986), Randall and Randall (1987), Schultz (1953a; 1953b; 1960; 1966), Melanesia Burgess and Axelrod (1975a) Micronesia Rofen (1961), Amesbury etal. (1979), Myers (1988; 1989), Randall (1960a) New Zealand Whitley (1968) New Caledonia Fourmanoir and Laboute (1976) Palau Helfman and Randall (1973), Matsuura (1982) Papua New Guinea Munro (1967), Kailola (1975; 1987a; 1987b) Polynesia Bagnise/a/. (1974) Samoa Jordon and Seale (1906), Fowler and Silverster(1922), Schultz (1943), Wass(1984) Solomon Islands Harry (1949) Tuamotu Harry (1953) Yap Amesbury (1978), Matsuura (1982) 210 Andrew Wright taxonomic reviews are continually being added to the literature. It is difficult for fisheries workers in the region to monitor continually new literature as it becomes available as personal contacts with the taxonomists involved and access to the literature is difficult for the majority of investigators involved in research at the national level. In excess of 300 species of finfish have been recorded from reef-associated fisheries throughout the Pacific. Although the taxonomy of most genera is relatively stable, the 50 or more families represented in tropical reef-associated fisheries are subject to regular review by taxonomists world-wide. Reviews and revisions of the taxonomic status of tropical food-fish families have regularly appeared in the scientific literature. The families presenting the most problems include those which exhibit colour change with age, those that are sexually dichromatic and those that change both sex and colour with age. The sequence of colour change in some species is associated with patterns of sexual ontogeny. Scarids (Randall, 1963), some lethrinids (Young and Martin, 1982), nemipterids (Young and Martin, 1985) and all serranids so far studied in detail (Shapiro, 1987) are protogynous hermaphrodites with most females eventually developing to males by sexual inversion. However, the proportion of primary males, primary females and transitional-sex individuals in a population varies between species and probably within a species, among locations. For the untrained, colour changes associated with sexual ontogeny create significant difficulties with species identification. Analysis of data for incorrectly identified species can result in wasted research effort; thus, after appropriate research programme design, accurate identification of species should be a priority for the execution of effective marine resource research in the South Pacific. Among the scarids and some of the balistids, for example, colour is an important character in distinguishing species and sex. However, the use of this as a diagnostic feature requires caution. Choat and Randall (1986) reviewed the scarid family, providing a much needed and authoritive guide to the scarid species from the Great Barrier Reef. This is generally applicable to the South and western Pacific although there are distinct regional groupings of scarids in some locations, e.g. the western Indian Ocean, the Indo-west and central Pacific (Randall and Bruce, 1983). Important recent reviews of other Indo-Pacific tropical fish families include the scombrids (Collette and Nauen, 1983; Collette and Russo, 1983), lutjanids (Allen, 1985, Allen and Talbot, 1986), and caesionids (Carpenter, 1987). In addition genera are also often reviewed. Recent examples include a review of Calotomus and Leptoscarus (Bruce and Randall, 1985),Plectropomus (Randall and Hoese, 1986) and Pinjalo (Randall et al, 1987). New technology has the potential to revolutionise the accessibility of information concerning fish species groups throughout the region. Most na- Shallow Water Reef-associated Finfish 211 tional fisheries administrations in the South Pacific have successfully adopted computers as an important tool to assist administration and research. The FAO and the International Center for Living Aquatic Resources Management (ICLARM) are collaborating to develop a computer-based system that will summarise information similar to that provided through the FAO species synopses. The objective is to develop a system, called FISHBASE, that in addition to providing biological and fisheries information, will incorporate a procedure facilitating species identification (Froese, 1990; Froese et ai, 1992). Once compiled and an operational system developed, this software will be available in a CD-ROM format and annually updated.

HABITAT AND FINFISH DIVERSITY

Coral reefs provide habitats for the most diverse offish communities found in marine environments. The habitats of the reef-associated finfish resources considered here are almost as diverse as the families of fish under consideration. Because of the variety of transition zones between habitats associated with coral reefs and the ability offish to transverse and inhabit more than one, environmental and habitat preferences for individual species, espe­ cially those targeted by fisheries in the Pacific islands, can often only be defined in broad terms. Throughout Melanesia and in some of the islands of Polynesia and Micro­ nesia, mangroves are found in close association with coral reefs. In many places they form contiguous habitats, with mangroves establishing themselves along the shore on consolidated reef platforms. Unlike the large deltaic swamplands associated with rivers in Papua New Guinea, supporting some of the largest coastal mangrove ecosystems in the world, the mangrove estuaries of Solomon Islands and islands to the east and north are relatively small and are isolated from each other by extensive coral reef lagoons (Stoddart, 1969). The fish fauna associated with mangroves systems throughout the Indo- Pacific region has been discussed elsewhere (Haines, 1979; Blaber, 1980; Collette, 1983b; Gunderman and Popper, 1984; Blaber et ai, 1985; Blaber etal., 1989; Blaber and Milton, 1990). Such systems have a characteristic fish fauna comprising permanent and temporary residents, the transients utilising the estuaries during feeding forays or on a longer term basis for a portion of their lives(Yanez-Arancibia^a/., 1980; Belle/ ai, 1984; Collette, 1983b; Kulbicki etal, 1992; Thollot, 1992). Snedaker (1978) estimated that up to 90 per cent of marine species in the tropical Indo-Pacific are found in mangrove systems during one or more periods of their lives. Collette (p. 95, 1983b) noted "the success of fisheries in many tropical regions may be closely tied to the health of the mangrove swamps which 212 Andrew Wright serve as important nursery grounds for species such as snappers (Lutjanidae), jacks (Carangidae), and mullets (Mugilidae)". This was supported to some degree by Robertson and Duke (1987) who noted the importance of mangrove systems in northern Australia as nursery sites for commercially important fish stocks. However, Parrish (1989) noted that although mangrove and seagrass communities appear to offer attractive habitats for juvenile fishes, there is no clear evidence that reefs situated favourably to shallow water vegetation habitats experience enhancement of early life stages through recruitment by comparison with more remote reefs. Recent studies suggest that few fish associated with coral reefs as adults utilise mangrove habitats during the juvenile stage of their life (Quinn and Kojis, 1985). Blaber and Milton (1990) reported that less than 10 per cent of juvenile fish species sampled in Solomon Islands were reef-associated species and that few of more than 500 species recorded from coral reefs in the area occurred in adjacent estuaries. Blaber and Milton (1990) recorded 136 species offish in 13 estuaries they sampled from mangrove systems in Solomon Islands noting that this was close to the number recorded from other tropical Indo-Pacific estuaries. The extent of utilisation of mangrove ecosystems by reef-associated fish requires further study. In particular, differences in species composition between sites subjectto different degrees of freshwater influence, warrant closer examination. Researchers have similarly debated the utilisation of seagrass communities by fish that spend the majority of their lives on coral reefs. Amesbury (1988) reviewed the physical relationships and biotic interactions among coastal habitats in the tropical Pacific and suggested thatmangrove habitats and seagrass beds do not serve as nursery areas for the majority of reef-associated fish. He concluded that juveniles and adults of tropical shorefishes tend to occur in the same habitat. However, Derbyshire and Dennis (1990) found that of fish sampled from seagrass habitats in northern Australia, a large portion were coral reef-associated. In their view, this confirmed that the nature of adjacent habitats did effect the faunal composition of seagrass communities. This supported earlier work by Heck and Weinstein (1989) in which feeding by juvenile lutjanids, scorpaenids, haemulids and mullids was found to be concen­ trated in seagrass meadows. The export of nutrients and organic matter transported from seagrass communities to reef environments by migrating fishes was studied by Meyer and Schultz (1985) who monitored the feeding ofjuvenil e haemulids over seagrass beds during the night and subsequent excretion of wastes over reef habitats during their daytime resting periods. There are few other examples of this type of study in the literature. Coral reefs are defined as carbonate structures at or near sea level which support viable populations of scleractinian corals (Preobrozhensky, 1977). The Shallow Water Reef-associated Finfish 213 extent of coral reef habitat in the South Pacific is yet to be accurately quantified. Smith (1978) estimated that the South Pacific region, including eastern Australia, supported 77,000 km2 of coral reef extending to a depth of 30 m and that this represented 13 per cent of the total global estimate for reef habitats. UNEP/IUCN (1988) describe present day reefs as falling in two basic categories: those established on continental shelves of larger land masses and those that develop adjacent to deep water in association with oceanic islands. Within these two categoriesare anumber of differentreef types: fringing reefs which grow close to shore (i.e. most shelf reefs, although some develop around oceanic islands); patch reefs which form on irregularities on shallow parts of the sea bed; bank reefs which occur deeper than patch reefs, both on the continental shelf and in oceanic waters; barrier reefs which develop along the edge of the continental shelf or through land subsidence in deeper water and are separated from the mainland or island by a relatively deep, wide lagoon; and atolls, which are roughly circular reefs around a central lagoon and are typically found in oceanic waters, probably corresponding to the fringing reefs of long since submerged islands (p. xvi; UNEP/IUCN, 1988). Information on the status of coral reefs in the South, central and western Pacific is constrained by the poor status of basic research on reef ecology throughout the region. To a large degree, generalisations on reef ecology are drawn from detailed research carried out at Enewetak Atoll in the Marshall Islands between 1954 and 1984 at the Mid-Pacific Research Laboratory under the auspices of the U.S. Department of Energy. With the discontinuation of research at Enewetak, coral reef-related research in the region is concentrated in French Polynesia and New Caledonia, on the Great Barrier Reef off the Australian east coast and to a lesser degree in Micronesia through the University of Guam and the Marine Science Institute in the Philippines. Dahl (1985) and UNEP/IUCN (1988) summarise available information on reef habitats through­ out the region. Increased destruction of coastal habitats and the implications for fisheries production have been discussed by a number of authors (Brackel, 1981; Perm, 1981;Dahl, 1985;Thorhaug, 1989; Hatcher^/., 1989;Wilkinson, 1992). As­ sessment of the impact of coastal habitat alteration resulting from natural or man- induced alterations is constrained by a general paucity of quantitative data that can be used to describe change. However, it appears that man does currently possess the knowledge and technology to mitigate impacts of many sources of man-induced stress (Grigg and Dollar, 1990). These authors argue that al­ though coral reefs ecosystems may be reasonably well adapted to absorb and respond to stress over the long term, as population growth and man's utilisation 214 Andrew Wright of reef resources continues to escalate, there is an increasingly urgent require­ ment to develop sound management practices for reef systems. They prescribe a need for a quantitative index for measuring stress to aid in the relative assessment of natural versus anthropogenic impact. Unless effective manage­ ment procedures are introduced and accepted in the near future, coral reef systems face extinction (Wilkinson, 1992). Theory concerning the relationship between habitat diversity and species complement has been summarised in many ecological texts (e.g. Krebs, 1972; Barnes and Hughes, 1982). Discussion of mechanisms governing the partition­ ing of available niches within coral reef-associated environments, commenced by Smith and Tyler (1972) and Russell et al. (1974), has resulted in the development of two theories, the "competition" or "order" hypothesis (Smith and Tyler, 1973; Watt, 1987) and the "lottery" or "chaos" hypothesis (reviewed by Bohnsack, 1983; Sale, 1980; 1982). Somewhat different from these was the antagonistic or critical unit hypothesis proposed by Bradbury (1977). The mechanisms influencing local diversity have been suggested to include niche specialisation (Smith and Tyler, 1973; Watt, 1987; Greenfield and Johnson, 1990), random colonisation determined by the availability of space (Robertson and Lassig, 1980; Williams, 1980; Bell and Galzin, 1984; Findley and Findley, 1985), habitat productivity and structure (Bohnsack and Talbot, 1980; Williams, 1982; Sale and Douglas, 1984; Bell and Galzin, 1984; Findley and Findley, 1985; Shpigel and Fishelson, 1986), competitive interactions (Sale etai, 1980; Gladfelter and Johnson, 1983; Shulman, 1985; Sweatman, 1985; Foster, 1985; Shpigel and Fishelson, 1986), variable recruitment patterns and predation (Talbot et al., 1978), and symbiotic-type dependencies (Ormond, 1980; Robertson and Polunin, 1981). The "evidence" supporting the various schools of thought relating to factors determining finfish diversity in coral reef habitats have been reviewed by Ehrlich (1975), Sale (1980), Sale and Williams (1982), Montgomery (1990), most recently by Sale (1991) and are discussed by Pyle (this volume). These studies and the resulting hypotheses relate principally to shallow water fish communities relatively well confined within close vicinity of a coral outcrop or bommie, a patch reef or within a well-defined area of reef slope. The extension of this theory to relatively transient families of demersal and neritic pelagic fishes contributing to island fisheries such as lethrinids, nemipterids, lutjanids, mugilids, mullids, carangids, scomberids, sphyraenids, and serranids (although the later are somewhat more site-attached), has yet to be attempted. However, recent studies involving trapping and tagging of reef- associated species {e.g. Dalzell and Aini, 1987; Davies, 1989; Newman, 1990; Whitelaw et al., 1991) have the potential to provide important information relating to recruitment and mortality, residence times, standing stock and migration among or within habitats. Shallow Water Reef-associated Finfish 215

FEEDING AND TROPHIC RELATIONSHIPS Fishermen throughout the South Pacific exploit information on the food habits of fishes to target preferred species from the large diversity available. This information has been accumulated over generations of affiliation with the reef- associated fish fauna as an important subsistence source of food. However, over the last four decades a significant amount of information on the diets and feeding habits of reef-associated fish has been published. Because of the large number of species offish taken by island fishermen and the huge diversity of food items consumed by these fish, a comprehensive review of the diets of reef-associated fishes is not attempted here. With respect to fish targeted by island fisheries, researchers are referred to Kow (1950), Hiatt and Strasburg (1960), Randall (1960b), Randall and Brock (1960), Hida (1972), Hobson (1974), Harmelin- Vivien and Bouchon (1976), Hobson and Chess (1978), Walker (1978), Randall (1980), Harmelin-Vivien (1981), Sano et al. (1984), Parrish et al. (1986), Parrish (1987), Blaber et al. (1990), and Blaber et al. (1990a; b) for examples of studies of the diets of reef-associated fishes and comparisons with diets reported for the same or related species elsewhere. Investigations of the food of tropical reef-associated fishes encounter difficulties associated with sampling sufficient numbers offish to ensure that a representative sample of the diet is examined. Few research programmes have been sufficiently well supported to undertake dietary studies for extended periods that are necessary to reveal dietary change that may be associated with reproductive cycles, changes in trophic relationships with size and age, depth or habitat differences with age or seasonal environmental change (see Helfman, 1978). Thus, information relating to individual species is often compiled from a number of studies which may be geographically separated. There are a number of significant constraints to studying the diets of tropical reef-associated fishes. The method of sampling is an important consideration as not all methods yield useful samples for subsequent examination in the laboratory. Spearing is perhaps the only method with which stomach contents can be assumed to provide a realistic indication of diet. If regurgitation occurs as a result of spearing, most larger items can be collected for subsequent identification (Randall, 1960b). However, fish that are trolled (e.g. Hida, 1972), handlined (especially those from deeper water), and even those netted or poisoned often regurgitate stomach contents (Treasurer, 1988). This results in a large number of empty stomachs which distorts subsequent analysis. Use of poisons also results in many of the larger piscivorous species consuming smaller species more readily affected by the poison (Hiatt and Strasburg, 1960). Traps usually only yield samples with evacuated stomachs or with stomachs contain­ ing prey ingested after entry to the trap. In addition to an accurate assessment of the representative value of the stomach sample as an indication of the diet, 216 Andrew Wright dietary studies in the South Pacific also suffer from a lack of taxonomic expertise available to identify food items. Dietary studies of fishes include those that attempt to determine the nutri­ tional relationship of one species with respect to other members of the commu­ nity thus determining competition and sharing of food resources (e.g. Staples, 1975; Oda and Parrish, 1981), and those that attempt to estimate the total amount of food consumed by a population offish (Hellawell and Abel, 1971; Eggers, 1977; Elliot and Persson, 1978; Majkowski and Waiwood, 1981). Windell (1968), Elliot (1979), Hyslop (1980), Persson (1986) and Jobling (1986) critically reviewed various methods including indices determined from fre­ quency of occurrence, volumetric or gravimetric composition in the diet and availability indices that link the abundance of prey in the environment to diet (e.g. Ivlev's electivity index [Ivlev, 1961]). Hyslop (1980) concluded that in order to describe diets accurately, at least one method that measures the quantity and another that measures the bulk of food material present is necessary. He noted the importance of considering differen­ tial rates of digestion and the desirability of linking measurements of bulk to stomach capacity. On the basis that graphical representation of data is relatively easy to interpret, Mohen and Sankaran (1988) developed two compound indices for dietary analysis. These procedures were used by Costello (1990) to develop another graphical method utilising the frequency of occurrence and the relative abundance of the prey in the diet. In addition, the model provides some indication of the relative importance of each prey and the degree of homogeneity of prey selection in the predator population. With respect to food fishes of interest to island fisheries, Randall and Brock (1960) simply noted, to the lowest taxonomic level possible, the diet of epinephelids and serranids in the Society Islands, measuring the lengths of identifiable food items. Hiatt and Strasburg (1960) and Shpigel and Fishelson (1989) listed the number offish species containing a particular item of food, thus providing information on the predilection of each fish species for particular items in the diet. Randall (1960b) measured the volume of each food item in the stomachs of fish he studied in the West Indies. Sana et al. (1984) applied methodology developed by Hobson (1974) to determine, 1) the percentage of samples containing a particular prey item, 2) the mean per cent of the item in the diet volume, and 3) a ranking index, which excluded samples with empty stomachs and which was obtained by multiplying the first and second values. Wahbeh and Ajiad (1985a) used a similar combination of measure­ ments (numerical, frequency of occurrence and gravimetric) to determine relative indices of importance for the goatfish, Parapeneus barberinus, in Jordan. Blaber (1986) also used a per cent frequency of occurrence of each prey and measured the selection of prey by predators using three indices of electivity, including that of Ivlev (1961). The frequency of occurrence and volume of each Shallow Water Reef-associated Finfish 217 prey item were also measured for a study of the diets of mullids in the north-west Hawaiian Islands and an Index of Relative Importance (after Pinkas et al., 1971), calculated (Sorden, 1983). Hobson and Chess (1978) recorded the number, size range, a subjective assessment of the state of digestion and a volumetric measurement of each food item for planktivorous fishes at Enewetak Atoll. Goeden (1978) applied a subjective measurement (after Hobson, 1968) to the stomach contents of Plectropomus leopardus collected from the Great Barrier Reef. Kow (1950) employed a less refined subjective measure during his study of the food of fishes of the Singapore Straits. Sainsbury and Whitelaw (1984) also visually estimated stomach fullness and the presence or absence of prey (to the taxonomic level of class or subclass) was noted. Brewer et al. (1989) and Blabere/a/. (1990a; b) sorted stomach contents by species. Each component was dried to constant weight at 60°C and the diets of each species were expressed in terms of the percentage contribution of each prey category to the total dry weight of stomach contents. Sainsbury (1986) described a method for estimating daily food ration from field observations of the diel cycle of stomach contents, without the need for independent determination of gastric evacuation rate. As part of a study to estimate the effect of feeding ofLutjanus russelli on commercially important penaeid prawns on the Queensland coast, Smith et al. (1991) fed rations of penaeid prawn and pilchard foods at different temperatures to determine the relationship between feeding and growth. Thresher and Colin (1986) described the trophic structure of the fish community of the reef slope at Enewetak Atoll on the basis of visual observations and known diets of the fish encountered. These examples, which cover most procedures so far employed in the study of the diets of tropical reef-associated fishes, are notable for their concentration on piscivory {e.g. Hobson, 1979; Diamant and Shpigel, 1985; Parrish et al., 1986; Parrish, 1987). Study of the diet of the herbivores and omni vores present additional difficulties, especially with respect to quantitative assessment of the proportion of different items in the diet such as diatoms, foraminifera and filamentous algae. In addition the assessment of the importance of non-target food items ingested incidentally, for example the importance of detrital particles and thin mucoid layers formed around substrate particles by blue-green algae and bacteria, is difficult to determine (e.g. see Jones, 1968; Robertson et al., 1979; Hatcher, 1981). Although the physical and biological features of coral reefs associated with geographical differences may affect the food habits of reef fish species among locations, most fish can be categorised as carnivorous, herbivorous or omnivo­ rous throughout their geographic range. Hiatt and Strasburg (1960) further partioned the fish community at Enewetak Atoll and identified four groups of herbivores: those primarily feeding on unicellular algae {e.g. some mugilids and blennids); grazers that crop algae close to the substrate (e.g. some leiognathids, 218 Andrew Wright siganids, acanthurids, labrids, gobiids, blennids, balistids, monocanthids, ostracionids, and canthigastids); browsers with teeth capable of pruning algal fronds (e.g. some siganids, the kyphosids, and some acanthurids, balistids and tetradontids); and the incidental herbivores where algae are ingested in associa­ tion with other primary foods (e.g. chaetodontids, pomacentrids, labrids, scarids, gobiids, and tetradontids). Among the relatively few detritus feeders they identified were some of the mugilids, gobiids and blennids. Of the 233 species offish that they examined (approximately one third of the total number offish species recorded from the Marshall Islands at the time of their study), Hiatt and Strasburg (1960) identified only one scavenger, the nurse shark. However, some of the lethrinids and nemipterids also fit this category (e.g. Aldonov and Druzhinin, 1978). Few species examined were classified as phytoplankton feeders but a number relied on zooplankton. These include the dussumierids, engraulids, atherinids, hemiramphids, and manta rays. In addition, some scombrids such as Rastrelliger kanagurta, at least one shallow water lutjanid, Macolor niger and the caesionids are zooplanktivores. Hiatt and Strasburg (1960) classified the carnivores into four broad groups, those that feed princi­ pally on fossorial fauna (resident within the substrate), benthic fauna, mid-water fauna and the roving carnivores that are either largely demersal and strongly reef-attached or those that are more pelagic in nature and frequent reefs to feed. Food fishes targeted by island fishermen classified as carnivores of fossorial fauna include sparids, holocentrids, polynemids, lutjanids, mullids, many of the labrids, balistids and ostracionids. Included among the carnivores of the benthic fauna are the carpet sharks, most of the congrid and muraenid eels, synodontids, polynemids, holocentrids, lutjanids, leiognathids, lethrinids, labrids, serranids, carangids, balistids, scorpaenids, sparids and mullids. Carcharhinids, belonids, holocentrids, fistularids, scombrids, carangids, lutjanids, and sphyraenids were described by Hiatt and Strasburg (1960) as including carnivorous species that feed principally on mid-water fauna. Demersal and pelagic carnivores include many of the fish species taken by South Pacific islands fisherfolk. The demersal grouping includes carcharhinid sharks, carangids, and serranids such as Plectropomus spp. Those identified as largely pelagic, transient roving carni­ vores include at least one lutjanid, Aprion virescens, additional carcharhinids, scombrids such as the dogtooth tuna, Gymnosarda unicolor, carangids such as Caranx ignobilis, Spanish mackerel, Scomberomorus commerson and the sphyraenids. Randall (1960b) developed a slightly different system for categorising the trophic relationships offish on reefs in the West Indies. He divided the 212 species he examined into seven groups based on food habits; plant and detritus feeders, zooplanktivores, "shelled"-invertebrate feeders, generalised carni­ vores, ectoparasite feeders, and fish feeders. Hobson (1974) analysed the stomachs of 1,547 fish representing 102 species from Hawaii and described a Shallow Water Reef-associated Finfish 219 different system of classifying reef-associated fishes on the basis of their trophic relationships. He divided generalised carnivores into those that are primarily nocturnal predators, crepuscular predators, and diurnal predators. Within these categories he identified those species that are specialised to ambush prey, those that stalk, those that hunt prey in crevices, those with sensory specialisations that detect concealed prey, predators capable of taking prey from among the plankton during the day and predators that prey on benthic invertebrates during the day. In addition, he noted that herbivorous fishes have many of the behavioural characteristics of the diurnal predators specialised to prey on benthic invertebrates. Sano et al. (1984) examined the stomachs of 1,891 specimens from 188 species taken from reefs in Okinawa. On the basis of their analysis they classified the food habits of the species investigated into eight broad categories, largely following the categorisation suggested by Randall (1960b): herbivores (divided into grazing and browsing herbivores), zooplankton-feeders (midwater and demersal zooplanktivores), benthonic ani­ mal-feeders (mobile and sessile benthonic prey forms), omnivores (principally algal, principally animal and generalised omnivores), coral polyp-feeders, ectoparasite feeders, fish skin-feeders and piscivores. Schooling as an effective mechanism employed by prey as a tool to evade predation and by piscivores to assist their predatory activities has been studied by a number of researchers (Davis and Birdsong, 1973; Neill and Cullen, 1974; Ehrlich, 1975; Major, 1977; Hobson, 1973; 1975; 1979; Murphy, 1980; Aoki, 1982; Hasegawa and Kabayashi, 1988). Researchers are referred to this litera­ ture for detailed accounts of the theory of schooling and the role of schooling in trophic interactions among the reef fish community. Despite the fact that the trophic particulars of many reef fish species have now been reasonably well described, community trophic relationships among South Pacific reef habitats remain relatively poorly understood (Parrish et al., 1985; Parrish and Dee, 1992). The description of the trophic relationships of fish communities on coral reefs was firstattempte d in detail by Odum and Odum (1955) in their classic study at Enewetak Atoll in which they described the trophic structure quantitatively as pyramids of mass. They ascertained that pyramids of biomass structure show up within some taxonomic groupings, citing fish as an example. They determined that there was a predominance of herbivores, e.g. scarids, acanthurids, pomacentrids and chaetodontids in com­ parison to the weight of labrids, serranids and other carnivores. However, other workers, for example Goldman and Talbot (1976), reported that piscivores may account for more than 50 per cent of the fish biomass (see below). Recently there have been attempts to incorporate quantitative information on trophic relationships into ecosystem box models (Andersen and Ursin, 1977; Laevastuand Larkins, 1981; Livingston, 1985; Polovina, 1984a). The ECOPATH model, for example, "is an analytical procedure to estimate the biomass budget 220 Andrew Wright for a box model of an ecosystem given inputs which specify the components of the ecosystem, together with their mortality, diet, and energetics value" (Polovina, 1985; p. 457). The ECOPATH model is not a dynamic simulation model with a time component; rather it estimates the biomass budget for a marine ecosystem in a static situation under the assumption that the ecosystem is at equilibrium conditions. In few situations has sufficient information been available to input to such models although Polovina (1984b) did apply data for the marine community at French Frigate Shoals and Alino et al. (1992) reported on similar work in the Philippines. Parrish et al. (1986) and Norris and Parrish (1988) studied the diets of populations of piscivores in a remote location in the Hawaiian Islands, estimated the standing stock of piscivores as a proportion of numbers and biomass of the total community of fishes and attempted to quantify the importance of the various piscivores and prey to the entire fish community. An important consid­ eration in any attempt to model community structure on coral reefs concerns mortality caused by the removal of components of the community, in many situations mainly the piscivores, through fishing, the parameter "F" in the Bulk Biomass Model (Livingston, 1980; 1985) and annual fishery catch data (Polovina, 1984a). Trophic relationships among the fish community targeted by reef fishermen can alter as a result of the selective removal of some components of the community (Munro and Williams, 1985; Russ, 1985; Munro et al., 1987). Al­ though the factors governing fish community structure are not well understood (for further discussion on the lack of data demonstrating the direct role of predation in determining the structure of reef fish assemblages see Sale, 1980), it has been assumed that an uneasy equilibrium does in effect exist and that prey species are controlled at a particular biomass by the combined effects of growth and recruitment on one hand and immigration and mortality on the other. An increase in the mortality caused by fishing will result in a new equilibrium at a lower biomass than that prior to fishing (Munro and Williams, 1985). The lower biomass equates directly to a decrease in catch per unit effort and on this basis, suggests that if the biomass of predators, such as sharks, is reduced, the biomass of prey species will increase (Pauly, 1979; Munro and Williams, 1985; Munro etai, 1987). Although it is widely recognised that fishing will affect the structure of reef fish communities, few studies have attempted to describe this effect quantita­ tively and many researchers have called for more work in this area (e.g. Sainsbury, 1982). As piscivores comprise up to 54 per cent of the biomass of the fish community of some reefs (Parrish et al., 1986) and are usually the target component of most reef-associated fisheries, it should be possible to describe change in community structure as a result of fishing. This was attempted in an experiment akin to applying fishing pressure on top level predators by Stimson Shallow Water Reef-associated Finfish 221 et al. (1982) who removed muraenid eels from a small patch reef and monitored the abundance and size of the remaining reef fish populations over one year. They concluded that the presence of predators such as eels is important in structuring reef fish communities. In a similar study, Shpigel and Fishelson (1991) removed all individuals of three species ofCephalopholis from a area of reef in the Gulf of Aqaba noting that after three years the initial population composition was still not restored and that the habitat was replenished by other predatory species of different taxa with only partial (21.6 per cent) replenish­ ment by species of the same genus. They noted that the removal of this component of the community resulted in little change in the abundance and species composition of the community typically preyed upon by Cephalopholis. The relationship between environmental parameters and preda­ tory behaviour of reef fish, a topic that has not received much attention in the literature, is discussed by Gyuris (1992). Randall (1982), Bohnsack et al. (1992) and Russ (1992) discussed the utility of marine reserves in reef fisheries management, Russ (1985) and McManus et al. (1992) described differences between reef areas subject to fishing and adjacent areas protected from fishing in the Philippines and Polunin and Roberts (1992) describe a similar situation in the Caribbean. Johannes (1977; 1978a; 1981a) describes the application of an understanding of the value of closed areas in traditional fishing practices throughout the Pacific and Polunin (1990) queried the value of marine reserves as effective mechanisms for resource conservation as applied through traditional systems in Indonesia and Papua New Guinea.

REPRODUCTION

The reproductive biology of reef fishes targeted by island fishermen is, in most cases, complex, with a wide variety of reproductive strategies exhibited. Much work remains to describe satisfactorily the life history of the majority offish species encountered in fish landings in the South Pacific. An attempt is made here only to introduce the topic, providing some indication for the major food-fish groups, of current knowledge on reproductive biology. Thresher (1984), Smith (1982) and Turner (1986) provide reviews of the reproductive biology of most reef fishes and readers are referred to those publications for more detail. In order to gain an accurate understanding of the reproductive biology of fishes, a random sample of the population is required. Thus, specimens for reproductive study collected from commercial catches should be supplemented with specimens collected by other means so that a representative sample of the total population is considered. Bagenal and Braum, (1968) and West (1990) have reviewed methods of collection, preservation, examination and interpre­ tation of results for research on the reproductive biology of fishes. 222 Andrew Wright

Reproductive activity in reef fishes is monitored either by direct observation of mating and spawning activity, for which there are many examples (e.g. Johannes, 1978b, 1981a, 1988; Robertsonsai, 1979; Robertson, 1983; Bell and Colin, 1986), or by monitoring gonadal development and relating this to other parameters such as age or growth (e.g. Fowler, 1991). Ovary develop­ ment is monitored macroscopically based on the external appearance of the ovary, and the oocytes within it, which can be broadly categorised into 5 to 8 stages (see Kesteven, 1960; Bagenal and Braum, 1968; Munro, 1976a; West, 1990). It can also be monitored histologically providing precise information on oocyte development. However, standardisation of terms to describe structures within the ovary, and the changes they undergo with development and recovery has not occurred, and this has led to some confusion (West, 1990). Monitoring the development of oocytes in the advanced mode microscopically is relatively simple. Besides providing information on size at maturity and breeding season it is also useful for estimating fecundity. West (1990) provides a detailed review of the assess­ ment of ovarian development in fishes. Some researchers have related the weight of the entire ovary to fish size to provide a measure of reproductive activity. Schaefer and Orange (1956) developed a gonad index (GI): GI = w/L\ 108 where w is the ovary weight (g) and L is fish length (mm). Although this, or variations of it, has been applied in numerous studies since, West (1990) cautioned that this may not provide an "index" that is independent offish size and that the relationship between gonad index and body size may be different for different stages of development. However, it may supplement macroscopic staging methods or oocyte measurements to provide greater clarity in describing reproductive activity. Most fishes have separate sexes (gonochores). This group is well repre­ sented in the catch of island fishermen in the South Pacific by holocentrids, mugilids, mullids, scombrids, haemulids, sphyraenids, gen-ids, siganids, belonids, hemirhamphids and carangids, among others. Many fishes associated with coral reefs are sequential hermaphrodites (change sex), while other species can be simultaneous hermaphrodites (displaying characteristics of both sexes at one time), which is common in deep sea species but for which there are few records from South Pacific shore fishes (Myers, 1989). Serranids (Reinboth, 1975) and lethrinids (Loubens, 1980) may be exceptions. Fishes that change from female to male, termed protogyny, include representatives from the scarids (Choat and Randall, 1986), serranids (Chen et ai, 1980; Warner, 1984; Shapiro, 1987), lethrinids (Young and Martin, 1982), nemipterids Young and Martin, 1985) and labrids (Robertson and Warner, 1978). Labrids and scarids are possibly the only Shallow Water Reef-associated Finfish 223 families among these in which primary males occur (Robertson and Warner, 1978; Warner and Robertson, 1978). Protandry, or sex change from male to female, is known among platycephahds (Okada, 1966; Fugii, 1970), sparids (Zohare/a/., 1978), gobiids (Lassig, 1977), and muraenids (Shen et al., 1979). Sexual dimorphism has been described for a number of reef fish, e.g. the balistids (Zama and Hattori, 1975; Matsuura, 1976; 1980) and acanthurids (Robertson, 1983). In addition, although dichromatism is common, colour alone is not always a reliable character to distinguish between sexes. Among some serranids (e.g. Yasuda et al., 1977) and many scarids, for example, complex colour pattern changes associated with sexual change complicates the study of the population biology (Randall and Choat, 1980). Descriptions of colour patterns associated with different stages of the life cycle of these fish have not been widely distributed in literature available to South Pacific fisheries workers, so information collected from fish landing sites in this region may often result in biological data being assigned to incorrect species. The recent review of scarid taxonomy by Choat and Randall (1986) addresses this problem for this group. There are usually three characteristic colour phases in the scarids, with the terminology by which to describe them being adopted at the International Symposium on Intersexuality in the Animal Kingdom in Mainz in 1974. The juvenile phase applies to young individuals in which the sex is largely indeter­ minate. The initial or primary phase (IP), characterised by dull red, brown or yellow hues with few distinguishing markings, is prevalent in subadults, mostly females and some types of males. Terminal phase (TP) fish are relatively brightly coloured with blues and greens predominating (Choat and Randall, 1986). However, colour patterns can change rapidly and this is often associated with reproductive behaviour (Robertson and Warner, 1978). Through a process of sexual inversion, most females assume a male identity, resulting in what are termed secondary males. The proportion of males that never change sex (primary males) varies among species. Even for individual species, the propor­ tion of males may vary between locations as a result of biotic or abiotic influences. Sex change may be socially controlled and the presence of second­ ary males can inhibit females from changing sex. On the other hand, if the ratio of terminal males to females becomes unfavourably balanced, the dominant female will change sex (Myers, 1989). Females and primary males (which are generally smaller than secondary males) typically display IP colour patterns. Larger males display the TP colouration which may be either second­ ary or primary derivation. Females may assume secondary male identity and TP colouration by synchronous sex and colour change and primary males with IP colour may change colour to achieve the TP colour phase (Choat and Randall, 1986). 224 Andrew Wright

Scarids have been observed to undergo spawning migrations within lagoons and to the outer reef slope (Randall and Randall, 1963; Yogo et al., 1980; Johannes, 1981a; Choat and Randall, 1986; Colin and Bell, 1991). Some species are diandric, forming schools and spawning in groups often after migration to specific sites, while others are monandric, at times being strongly site-attached with haremic, pair spawning (Choat and Randall, 1986). Although the diversity of spawning behaviour exhibited by scarids means it is not possible to describe a generalised pattern applicable to the group, at least some members of the family are reported to spawn on a lunar rhythm (Johannes, 1978b; 1981a). Particular attention to the mechanisms influencing the evolution of protogyny and the biotic and abiotic factors affecting its development in the life of serranids was reviewed by Shapiro (1987). Sadovy and Shapiro (1987) and Shapiro (1987) were concerned at the proliferation of criteria for the identification of protogyny and listed ten features that can assist in its identification in fishes. They noted bimodal size- and age-frequency distributions, female-biased sex ratios (including size- or age-specific sex ratios), yellow bodies that are not clearly early stages of oocytic artresia, and oocytes in testesare nonspecific features that may have many causes, only one of which is protogyny. On this basis, Shapiro (1987) concluded that the literature contained only reasonable evidence for protogyny for 12 serranids and that statements that all groupers are protogynous (Smith, 1965; 1975) may be erroneous. However, he also noted that all groupers that have been carefully studied are protogynous. The relationship between age and sex change in the serranids is not yet well understood due to difficulties in determining age (Chen et al., 1980). If sex reversal is linked to age or size and is endogenously controlled, removal of large individuals could lead to a detrimental decline in the proportion of males in a protogynous, monandric population (Russ, 1991). Russ (ibid.) cited the obser­ vations of Thompson and Munro (1974) for serranids taken in a Jamaican trap fishery to support this. However, Russ (ibid.) also noted that Reeson (1975) could find no evidence for the effect of fishing on sex ratio among scarids on Jamaican reefs. The age at which mature females first appear in serranid populations varies significantly between populations studied and Shapiro (1987) concluded that there is not a characteristic size at which transition occurs and that change may be influenced by a combination of behavioral and environmental parameters. Shapiro (1987) noted that many previous studies of the sex ratio in serranid populations reported a predominance of females in the population. However, because many reports have relied on catches from commercial fisheries, with unknown biases for selectivity, the structure of serranid populations in the wild is not accurately known (Shapiro, 1987). Tran­ sitional fish have been present throughout the year in most populations studied. Serranids are widely reported to aggregate at seasonally predictable loca- Shallow Water Reef-associated Finfish 225 tions for periods of 1 -2 weeks to spawn {e.g. Johannes, 1978b, 1981a; Colin, 1982; Shapiro, 1987). At these times, females release between 0.5-5 million eggs according to Shapiro (1987), although Thompson and Munro (1978) reported that the ovaries of four species of serranid in the Caribbean each contained about 160,000 eggs. Lethrinids are another protogynous family important in the catches of Pacific islands fishermen. Loubens (1980) reported size-related differences in the ratio of males to females in six species from New Caledonia and suggested that for Lethrinus chrysostomus, L. lentjan and L. variegatus this may be due to sequential hermaphrodism. Young and Martin (1982) studied the reproductive biology of eight species of lethrinid from northern Australia. They concluded that the evidence for protogynous hermaphrodism was substantial and that, within the populations sampled, sexual transformation occurred over a wide range of sizes. These authors noted more research on mechanisms involved in sex change in lethrinids was required. Evidence for sequential hermaphrodism was also found in at least three species of nemipterid fishes in northern Australia (Scolopsis monogramma, S. taeniopterus and S. bilineatus), with male gonad morphology in S. peronii and Pentapodus porosus suggesting similar transition (Young and Martin, 1985). Among other families, the lutjanids appear to be gonochoristic throughout life although the population sex ratio as well as the sex ratio at size may frequently be skewed resulting from differential growth and mortality rates among the sexes (Grimes, 1987). Grimes (1987) reviewed the reproductive biology of the lutjanids, reporting that length at maturity as a proportion of the maximum length attained by individual species varied between species and that populations associated with islands matured at a relatively larger size than those associated with continents. He postulated that these habitat-associated differ­ ences may be related to available food resources. He also noted that continental populations and species exhibit extended spawning during the warmest months while insular populations and species spawn year round with two peaks annually. In Papua New Guinea peak spawning among many fish species occurs in the months around September and April, with the major peak around September, although at any period of the year, some individuals in fish populations appear to be sexually active (A. Mobiha, Fisheries Biologist, Department of Fisheries and Marine Resources, Papua New Guinea, pers. comm.). Spawning in lutjanids appears to be associated with the lunar cycle (Randall and Brock, 1960; Mizenko, 1984) although the number of egg batches produced each spawning has not been conclusively determined (Grimes, 1987). Al­ though there are problems associated with accurately estimating batch and egg fecundity, lutjanids are highly fecund, with large females producing 5-7 million 226 Andrew Wright ova (Grimes, 1987). The egg production process and its relation to body size, fecundity estimation, and realised fertility require thorough study (Grimes, 1987). Mullids are also gonochores and are relatively easy to sex by visual examination (Wahbeh and Ajiad, 1985b). They are reported to be mature by the first breeding season after spawning. In Hawaii this occurs between December and July (Moffit, unpubl. m.s.1). Differential growth rates of the sexes is apparent in some species (Munro, 1976a) and many species aggregate to spawn (Helfrich and Allen, 1975). Breeding among mugilids in Hawaii appears to be concentrated during the austral summer. The sexes are separate and spawning behavior has been observed for Crenimugil crenilabis at Enewetak Atoll (Helfrich and Allen, 1975). Grant and Spain (1975a and b) have reported in detail on the reproduc­ tive biology of other species of mullet widely distributed throughout the Indo- Pacific, Liza vaigiensis and Valamugil seheli and Silva and De Silva (1981) described the reproductive biology of the estuarine species, Mugil cephalus. The reproductive biology of a number of acanthurids has been described by Robertson et al. (1979), Robertson (1983), Fishelson et al. (1987) and re­ viewed by Thresher (1984). These studies report on pair formation, territorial defence of feeding areas, migrations to the reef edge adjacent to deepwater to spawn, spawning rhythms associated with the tide and lunar phase, and colour changes with respect to sexual and social behavior. In some species of acanthurids males may grow larger than females (macroandry), in others the sexes may attain similar sizes (isomorphism) while in other species, females may attain greater maximum sizes than males (macrogyny) (Robertson, 1985). Some species of acanthurid spawn as pairs; others aggregate in dense schools (Dalzell, 1989). Dalzell (1989) studied the reproductive biology of acanthurids in northern Papua New Guinea where he observed protracted spawning with seasonal peaks, the stimulus for which was ascribed to changes in environmental conditions. For six species of acanthurid he monitored in Kavieng Harbour, Dalzell (1989) recorded lengths at maturity to range between 50 and 70 per cent of the maximum length attained by each species. Robertson (1983) suggested that a combination of primary and secondary tidal rhythms may influence lunar spawning cycles in the acanthurids and Thresher (1984) noted that acanthurid species may spawn throughout the lunar month, particularly where the species concerned are pair spawners, not group spawners. The reproductive biology of many siganids is relatively well understood and the application of this knowledge to hatchery rearing has been attempted in Fiji (Popper et al., 1976) and Palau. Schooling and lunar spawning behavior among some siganids has been described by Tsuda and Bryan (1973) and Johannes (1978b, 1981a). Gundermannefa/. (1983)dividedthesiganidsintotwogroups on the basis of habitat, behavioral characteristics and colouration. One group Shallow Water Reef-associated Finfish 111 includes species that live in pairs, have limited home-ranges on coral reefs and are brightly coloured. This group isrepresentedby Siganuscorallinus,S. puellus and Lo vulpinnus. The remaining group school, at least at some stage during their life cycle, assume a colouration similar to that predominating in their frequented habitats, and may undertake substantial migrations. Examples in this group are S. rivulatus and S. canaliculatus. The eggs and larvae of most siganid species are demersal and sticky. The exception is the floating egg of S. argenteus possibly explaining the wide distribution of this species (Gundermaim et al., 1983). There are also a number of important pelagic fishes that are regularly taken by fishermen operating in oceanic waters close to coral reefs in the Pacific islands. These include the dolphin-fishes Coryphaena spp., the biology of which was reviewed by Palko et al. (1982), wahoo, Acanthocybium solandri, for which little information is available, the scombrids, Euthyunnus affinis and Auxis thazard, the biology for which was summarised by Yoshida (1979), Spanish mackerel, Scomberomorus spp. (see McPherson, 1981), and barracu­ das or sphyraenids. The biology of the Spanish mackerel, Scomberomorus commerson, has been reasonably well studied because of its commercial value (for a synopsis of Scomberomorus, see Collette and Russo, 1983). Four other species are found in the area of Papua New Guinea and Australia, with S. commerson alone extending its range as far east as Fiji. This species can undertake extended migrations, for example, along the Queensland coast where spawning occurs around the northern cays of the Great Barrier Reef in October and November. In other areas, for example in northern Papua New Guinea where some fish are resident year round, spawning occurs between July and November. Lewis et al. (1983) reported that Spanish mackerel spawn in Fiji between October and February with a peak in December. These authors also reported on the size distribution, sex ratio, age at maturity, age and growth for this species. Genetic analysis of liver and muscle tissue samples showed that the Fij i Spanish mackerel population is genetically distinct from northern Australian populations. Lewis et al. (1983) also reported on the biology of other scombrids, the double-lined mackerel, Grammatorcynus bicarinatus, dogtooth tuna, Gymnosarda unicolor and the mackerel tuna, Euthyunnus affinis. They also provided information for some sphyraenids and carangids. Although some species have been studied in detail, e.g. rainbow runner which is reported to spawn year round (Okiyama, 1970; Yesaki, 1979), little has been published on the reproductive biology of carangids. This is despite the fact the group contains a number of important food fishes (Johannes, 1981 a). Lewis et al. (1983) reported on size at maturity, sex ratios and spawning patterns for three species in Fiji {Caranx ignobilis, C. papuensis and C. melampygus), with notes on two others (Carangoides plagiotaenia and Caranx 228 Andrew Wright sexfaciatus). Johannes (1978b, 1981a) presented information on the spawning migrations of seven species of carangid from Palau, with indications of spawn­ ing activity relative to the lunar cycle. The linkage between lunar periodicity and spawning has been discussed widely inthe literature (e.g. Korringa, 1947; Johannes, 1978b, 198 la; Robertson, 1983; Taylor, 1984; Bell and Colin, 1986; Colin and Bell, 1991). Implications of fishing aggregations of spawning fish when they are particularly vulnerable to fishing have been discussed by Johannes (1978b, 1981a, 1988), Shapiro (1987) and Russ (1991). The effect of fishing targeting spawning aggregations, especially with respect to reproductive potential, requires significantly more research. In conclusion, studies by Warner (1975), Robertson and Warner (1978) and Shapiro (1983), cited in Montgomery (1990), demonstrate that reproductive biology of reef fish populations (age at reproduction, sex change or colour change) are influenced by environmental parameters, population structure, body size and behavioural interactions and that these aspects require careful consideration in life history studies.

RECRUITMENT

Recruitment has been described by Russ (1991, p. 614) and Doherty and Williams (1988) as "increases in the abundance of young offish produced by larval settlement". With respect to fisheries development and management, the definition by Gulland (1983, p. 117) is more applicable:".. .the process whereby young fish, previously inaccessible to the ordinary fishing gear become as a result of growth, change of behaviour or of movement on to the fishing grounds, potentially vulnerable to fishing". Almost all reef-associated fishes exploited by subsistence and commercial fisheries in the South Pacific produce pelagic eggs (Sale, 1980). The few exceptions to this include the majority of siganids, mentioned above, tetradontids and balistids, which nest (Johannes, 1981a). Among these families, eggs hatch within a day or two and the larvae become pelagic for about a month (Popper et al, 1976). The fate of eggs and newly hatched larvae of the demersal egg-layers after release to the water column has come under increasing attention during the last decade but the biology and ecology of juvenile reef fishes is still relatively poorly understood (Lobel and Robinson, 1986; 1988; Montogomery, 1990; Leis, 1992). The publication of guides for the identification of coral reef fish larvae, such as that of Leis and Rennis (1983) which covers 49 families, has enhanced the potential for improved research in this field. Spawning among many reef fishes often involves a migration, either to a reef location contiguous to oceanic water (e.g. Munro, 1976; Johannes, 1978b; Shallow Water Reef-associated Finfish 229

1981a for mullids, serranids, lethrinids and lutjanids), vertically in the water column {e.g. Helfrich and Allen, 1975; Bell and Colin, 1986 for mullids and caesionids respectively) or inshore {e.g. Lowell, 1971 in Johannes, 1978b for polynemids; Dekhnik et al., 1966 in Johannes, 1978b for sparids and Tiews et al, 1968 and Watson and Leis, 1974 for some of the small schooling pelagic species such as the engraulids and clupeids). Leis and Reader (1991) reported that milkfish {Chanos chanos) migrate up to 50 km to spawn on the Great Barrier Reef. Possible explanations for the evolution of these migrations include selection for behaviour which minimizes egg and larval predation by other reef residents (Johannes, 1978; Connell and Kingsford, 1992), promotes dispersal (Barlow, 1981), improves the likelihood of survival of pelagic larvae in waters with patchy and uncertain distribution of food (Doherty et al, 1985), reduces opportunities for predation on the spawning adults (Colin and Clavijo, 1978), or promotes fertilisation by simplifying the task of widely spaced male and female fish encountering each other (Moyer and Zaiser, 1981). Shapiro et al. (1988) noted the urgent need for comprehensive studies of spawning times and sites, and non-spawning times and sites with respect to current regimes prevalent at those locations and times. They drew attention to the need to consider relative scales with respect to distances and the sizes of fishes, i.e. at what scale should spawning site selection in response to current patterns be detectable: an island, reef system, individual reef or coral head? Similarly, Johannes (1978b) suggested study of the relative strengths of prevail­ ing currents during years of high and low recruitment could provide useful information on spawning strategy. Investigating the assertions of Barlow (1981) that larval dispersal is highly developed in reef fishes because reef fishliv e in a patchy environment and adults cannot migrate among patches and that reef instability has been an important factor selecting for dispersal, Doherty et al. (1985) developed a computer model to investigate the selective advantage to coral reef fishes of dispersal of sibling offspring. Colin and Bell (1991) examined the influence of tidal currents on the time of spawning and behaviour among 32 species of labrid and 13 species of scarid fishes at Enewetak Atoll noting that there did not appear to exist any generalised lunar spawning seasonality among labroids at this site. They suggested that lack of seasonality at Enewetak was because of the lack of environmental variability throughout the year at this site, although it was also noted, that some labroids will shift their spawning style under different current and tidal regimes. They also considered, that despite the presence of relatively more potential predators, the role of lagoons as nursery grounds for reef fishes required further research. In this respect, the issue of initial colonisation for species resident in isolated oceanic reef systems will also need to be addressed. 230 Andrew Wright

Russell et al. (1977) found substantial evidence for strong seasonal repro­ ductive activity for 54 species of coral reef fishes on the Great Barrier Reef. He noted that, although recruitment is strongly seasonal, there was little constancy in annual recruitment with high variability between years. Protracted spawning seasons and repeated spawning during the period of reproductive activity have been suggested as one mechanism for ensuring the wide dispersal of larvae (Sale, 1980). However, factors influencing dispersal of larvae and their subsequent recruitment to reef systems remain vague (but see Dufour and Galzin, 1992). Munro and Williams (1985) noted difficulties associated with tracking pelagic larvae, thus the source of subsequent recruit­ ment to reefs, were dependent on the duration of larval life, current strength and direction, swimming behaviour and larval mortality rates. They concluded that, although the extent of recruitment back to natal reefs remains undescribed, evidence for its occurrence is good. Some degree of self-recruitment, either from within lagoons or from the ocean, is essential among isolated, mid-ocean atoll systems in order for reef fish populations to be maintained. The decreasing diversity of reef species across the central Pacific to west longitudes in the South Pacific does suggest the length of larval life for many species is insufficient to survive dispersal in prevailing current systems. While Johannes (1978b) suggested many reef fish employ reproductive strategies favouring dispersal from the natal reef, he also discussed the possible return of larvae to islands or atolls after a suitable period as residents in gyres or eddies found in the lee of most island systems. However, although Bell and Colin (1991) considered that currents and eddies existed in the lee of Enewetak Atoll, they suggested that the majority of eggs, embryos and larvae exiting the lagoon would be lost to the atoll. Leis et al. (1991) examined the relative abundance of tuna larvae within 10 m of the surface in the lee of three islands in French Polynesia. They encountered high concentrations of scombrid larvae within 200 m of the reef, noting the majority of the larvae were in early stages of development. They were thus unable to comment on the duration of residence times in the nearshore environment but did suggest that the high productivity of nearshore environments offered significant advantages for the survival and development of tuna larvae relative to oceanic environments. Reef fish larvae may experience similar benefits in this habitat. Passive return in prevailing currents has been assumed to be the principal mechanism for subsequent location of larvae in an environment close to the reef immediately prior to recruitment (Leis and Miller, 1976). This has been suggested on the basis that larvae appear to be inadequately equipped to move against currents and because they are unlikely to have cues for the presence of reefs which could direct them until they were quite close by (Sale, 1980). How- Shallow Water Reef-associated Finfish 231 ever, Leis and Reader (1991) report that larvae of milkfish probably maintain a position vertically in the water column that promotes exposure to wind-driven currents to move inshore from offshore spawning sites on the Great Barrier Reef. The relationship between larval size and recruitment mechanisms in fishes, based on data from 72 species of marine and freshwater fishes was discussed by Miller et al. (1988). There are numerous records of the occurrence of larvae of reef fish offshore (e.g. Lobel and Robinson, 1988) but as noted by Munro and Williams (1985), there are many difficulties associated with securing a comprehensive knowl­ edge of pelagic larval life. Information from plankton tows in Hawaii have indicated that the larvae of reef species with pelagic eggs are more common 3 km from shore than closer (Watson and Leis, 1974; Leis and Miller, 1976) although goatfish larvae abundance peaks 0.5-2 kilometres from shore (Leis and Miller, 1976). Juvenile goatfish have been reported from the stomachs of tuna captured up to 16 kilometres offshore (Hunter, 1967). Studies on the genetic structure of populations suggest that only the most isolated will be totally independent of recruitment from other sources (Williams, 1983; Munro and Williams, 1985; Doherty and Williams, 1988). However, the observed lack of genetic differentiation in the few reef fish species studied is consistent with extensive dispersal but does not prove it (Munro and Williams, 1985). This aspect of reef fish biology requires significantly more research, including electrophoretic analysis, for an understanding of the distribution and maintenance of reef fish populations to improve. A final issue in this section concerns the impact of fishing on recruitment to reef fish populations. Russ (1991) noted that if the majority of reefs are not self- recruiting, i.e. they receive recruits from other reef systems up current, the relationship between stock and recruitment may be undetectable, and that even at a scale encompassing the range of any species of reef fish, although a relationship may exist, it is likely to be extremely variable in nature. In the South Pacific, Johannes (1978b; 1981a) has documented a widespread practice by fishermen to target spawning aggregations of reef fishes (Johannes, 1988). Russ (ibid.) was concerned that should no controls over this be exercised, stock sizes could be reduced rapidly, potentially affecting recruitment. Munro (1983) presented evidence to justify such concerns from the Caribbean. He noted the removal of significant numbers of adults appeared to affect spawning biomass leading to significant reductions in catch rates of targeted species among the lutjanids, serranids, haemulids, sciaenids, carangids and acanthurids, despite increased fishingeffort . Monitoring of the biomass of the spawning populations of deepwater fish in Hawaii is one mechanism employed in fishery management (see Somerton and Kobayashi, 1990). In that location overfishing is assumed to occur when the spawning stock biomass is 20 per cent of the virgin biomass. 232 Andrew Wright

GROWTH

Growth and mortality ofpost-settlemen t reef fishes has not received the same degree of attention in the literature as have the mechanisms governing recruit­ ment and reproduction. There is more information available on the apparent growth offish after recruitment to a fishery. The literature for these two subjects is growing extremely rapidly and is largely based on relatively recent techno­ logical developments with respect to electron microscopy for the study of hard parts, such as otoliths, in juvenile and adult fish and the widespread availability of computers and associated software to assist in the analysis of large data sets and complex questions relating to the analysis of length frequency data. Recognising that the timing and extent of settlement from the larval stage, is probably more important than spawning activity among adults in determining the initial sizes of successive age classes of fish populations on reefs (Sale, 1985), recent research to determine the history of post-settled larvae and juveniles has shown potential to expand our understanding of reef fish larval biology, especially with respect to ageing. There are three methods available for the estimation of the age in reef fishes, all of which have a number of assumptions associated with them. The methods include: 1) interpretation of rhythmic patterns on, and gross morphology of, hard parts (otoliths, bones and scales) as records of growth 2) analysis of the progression of modes in pooled length data for fish sampled from a population on the assumption that this progression records growth, and 3) by marking individual fish of known length for subsequent recapture and measurement with the assumption that any increase in length of individual fish between marking and recapture is attributable to growth. Chemical staining of hard parts is one method of marking and has most often been employed for later examination of otoliths to interpret growth (e.g. Dee and Radtke, 1989). Utilisation of spines and vertebrae for interpretation of growth have been reported by Lai and Liu (1979), Johnson and Saloman (1984), Edwards (1985), and Johnson (1985). Although scales are easy to collect and handle relative to otoliths, the advantage of otoliths lies in the relatively clear presence of apparent daily records of deposition. Although such records are also present in scales, their extreme thinness and their angular relationship with structural features means that interpretation is difficult (Pannella, 1974; Reynolds and Moore, 1982). A comparison between scale and otolith ageing methods in a species of herring is presented by Libby (1985) who concluded greater accuracy was obtained by otolith reading techniques in this species. Although some studies in the South Pacific region have interpreted ring deposition in otoliths as an annual event (e.g. Loubens, 1978; Chen etal, 1984; Uchiyama et ai, 1986; Baillon and Kulbicki, 1988; Ralston and Williams, Shallow Water Reef-associated Finfish 233

1988; Morales-Nin and Ralston, 1990), the majority have examined daily patterns. Daily patterns have been interpreted in otoliths for some time (Panella, 1971; 1974) to reveal age, spawning periods and major life history events such as recruitment (see Meekan, 1992). Subsequent research has substantiated the use of rhythmic patterns in otoliths as accurate indicators of age in juveniles, noting that as fish age, patterns become more difficult to discern and interpret as a result of overlayering or compression of patterns (Pannella, 1974; Brothers etal, 1976; Pannella, 1980; Campana and Neilson, 1985; Radtke, 1985,1988; Jones, 1986). Ring formation commences early in the life of most larval fishes, for example, in Pacific herring at the age of complete yolk absorption (McGurk, 1984). In most cases, patterns studied have been assumed to be daily and morphometric measurements of all three otoliths (sagitta, lapillus and asteriscus), have been related to fish size and possibly individual growth (e.g. Gjosaeter et al., 1984; Brothers et al, 1983; Radkte, 1985). Support for this has increased with the indication that deposition of growth increments appear to be controlled by neurosecretory proteins leading to an obligatory diurnal rhythm in otolith growth (Campana, 1984a, Gauldie and Nelson, 1988). However, increments formed as a result of circadian rhythm (once per 24 hr) can be induced through different processes from those induced through the action of environmental cues (often greater than once every 24 hr), thus explaining apparent morphological differences in increment structure noted in some studies (Campana and Neilson, 1985). Daily patterns have also been described for scales but not other hard parts. Assuming that otolith and somatic growth are related, incremental widths in otoliths should provide useful information on both size-at-age and instantaneous growth rates over short periods of time (Thorrold and Williams, 1989). In a study of a tropical sardine on the Great Barrier Reef, these authors found that cohorts separated by short periods of time may exhibit different growth trajectories. They noted that evidence was accumulating that indicates mortal­ ity rates of larval fish may be affected over relatively small spatial scales (1-100 m), and over temporal scales of hours to days. They cited the work of Fortier and Leggett (1985), Hewitt et al. (1985) and Taggart and Leggett (1987) to support this noting that factors affecting larval survivorship may be expected to vary on a similar scale (Legendre and Demers, 1984). As otoliths are not extensively remineralised in fishes, information about the physiological condition ofth e fish at the time of otolith deposition is retained. This can be interrogated to provide details on growth, nutrition and temperature for the duration of the fish's life (Struhsaker and Uchiyama, 1976; Brothers and McFarland, 1981; Campana, 1984a; 1984b; Radkte, 1988). Based on the temperature related exchange between strontium and calcium in otoliths measured by use of electron microprobe techniques, Radtke (1984; 1985; 1987; 1988) attempted to relate changes in otolith microstructure to 234 Andrew Wright physiological and environmental change. Thus, for example, Radtke (1985; 198 8) suggests the age at which the fish leaves the plankton and is recruited can be determined with daily resolution. Several studies have described otolith growth under different temperature regimes and food levels (e.g. Brothers, 1981; Brothers and MacFarland, 1981; Neilson and Geen, 1985; Campana, 1984a; Rice et ai, 1985). The majority of otolith studies for tropical reef fishes concentrate on juvenile life stages (Ralston and Miyamoto, 1981). This is because of the difficulty in interpreting marks in older fishes due to compression and compacting of marks. In addition, it is possible that the one-to-one relationship between increments and days is not maintained in older fish due to interruptions in growth (Pannella, 1971). In developing a methodology to age a deepwater lutjanid in Hawaii, Ralston and Miyamoto (1981; 1983) determined that within the sample examined, otolith length was allometrically related to fork length and that interruptions in daily ring deposition commenced at about the length fish attain maturity. The methodology involved examination of patterns of daily ring deposition in different sections of otoliths along a predefined axis and extrapo­ lation of this information to the total length of the otolith. This methodology has since been applied to a number of tropical species to estimate growth (Ralston and Miyamoto, 1981;1983; Morales-Nin and Ralston, 1990). Other studies relating otolith weight to growth have also been reported in the literature but Pawson (1990) notes that there is limited application of this methodology to fish populations with highly variable growth rates. Confirmation of growth estimates by a variety of techniques is desired where possible. In some instances, validation has been attempted using a combination of length frequency modal analysis and daily growth increments from otoliths (e.g. Ralston and Williams, 1988), periodic marks from dorsal spines and vertebrae (e.g. Johnson, 1985), increments on scales and otoliths (e.g. Libby, 1985), comparison between apparent daily and annual marks on the same otolith (e.g. Baillon and Kulbicki, 1988), chemical labelling (Ferreira, 1992), and tag- recapture information in conjunction with estimates obtained from marks on dorsal spines and otoliths (Prince et ai, 1986) or scales (Davis and Kirkwood, 1984). In a survey of 500 studies published between 1907 and 1980, Beamish and McFarlane (1983) reported that in 35 per cent of growth studies reported, no attempt was made to validate ageing techniques. They cautioned against the use of unvalidated age information and the errors that can result in understanding the biology and population dynamics. The costs of mistakes associated with errors in age estimation in management of commercial fisheries could be significant. Beamish and MacFarlane (1983) noted that only in mark-recapture studies or use of known age fish can all age classes in a population be validated. Examples of the use of tag-recapture methodologies to obtain information on Shallow Water Reef-associated Finfish 235 the growth of tropical reef fishes are not well represented in the literature. This is despite the fact that some of the earliest literature of coral reef fishbiolog y and ecology involved tagging. In 1958, Bardach tagged reef fish in the Caribbean to provide information on fish movements and in 1962, Randall tagged more than 4,000 fish captured in wire-mesh traps in the Virgin Islands to study movements and growth. In his report of the study, Randall (1962) provided information for 1,247 recaptures obtained over a 16 month period. He also included discussion on issues that effect the success of tagging studies such as tag-induced mortality, tag shedding, and apparent shrinkage. There are a number of assumptions associated with tagging (Gulland, 1983; Pauly, 1984). These include: 1) the natural mortality and vulnerability to fishing gears of tagged and untagged individuals is identical, 2) the tagged fish are randomly distributed in the population, 3) no tags are lost, 4) emigration and immigration are negligible, and 5) all recaptured fish are examined. The fact that tagging experiments have not often been attempted with reef fish populations is indicative that it is difficult to meet all these assumptions. The first assumption is difficult to assess although there are indications that mortality can be enhanced as a result of stress associated with the actual tagging operation (Randall, 1962; Ricker, 1975; Whitlaw and Sainsbury, 1986; Dekker, 1989). This can perhaps be reduced by use of a detachable tag system as proposed by Grimes et al. (1983) although information on growth will not be obtained by this method due to the lack of information on the length of the fish at the time of tagging. The second assumption is reasonable and tagged individuals of Acanthurus xanthopterus, A. nigricans, Scarus ghobban, and Caranx melampygus have been observed freely mixing in schools with untagged fish of the same species in northern Papua New Guinea (pers. obs.). However, 4-6 weeks after tagging, tags were observed to become heavily fouled with marine algae. The effect of this on the behaviour offish was not apparent but some recaptured fish with heavily fouled tags exhibited severe infection in dermal tissue surrounding the point of tag insertion. The third assumption is not possible to quantify so the challenge is to estimate the rate of retention. This can be attempted by double tagging. Whitelaw and Sainsbury (1986) noted that a combination of tag loss and tag-induced mortality in a tropical lutjanid gave a 70 per cent tag loss for animals tagged with dart tags and a 42 per cent loss for animals tagged with anchor tags. If conducted over a relatively short period of time (2-3 months for species targeted by tropical fisheries), migration may be assumed to be negligible. However, as noted above, many tropical reef fishes spawn throughout the year. If this produces recruits to the reef system through­ out the year, this assumption will not be valid. With respect to assumption 5), in the Pacific islands this can be reasonably easy to achieve. With effort expended on promoting public relations within the fishing community, including an explanation of the objectives of the project, and with perhaps the addition of 236 Andrew Wright an incentive to encourage returns offish with tags still attached, it is possible to gain access to a very high proportion of recaptures. Hilborn (1988) has examined this problem, mainly from the point of view of tags returned in commercial fisheries. Methods for the analysis of tag data for growth, on the assumption that growth can be described by a von Bertalanffy equation, are reasonably well described in the literature (e.g. see Walford, 1946; Beverton and Holt, 1957; Baker et al., 1991). For data involving animals at liberty for varying time intervals, Gulland and Holt (1959), Fabens (1965) and Munro (1982a) describe procedures for analysis. Chien and Condrey (1987) discuss biases in estimating growth param­ eters using Fabens' mark-recapture procedure. The Gulland and Holt procedure involves a plot of apparent growth against the mean size offish at tagging and recapture. This does not provide reasonable parameter estimates when the size difference between tagging and recapture is small or when the tagging procedure stunts growth. In such cases, a trial value of LM together with the mean of all mean lengths of returned fish and the mean growth increment can be used to estimate K, the Brody growth coefficient where: K = mean growth increment/L^- mean length. This is termed the "forced" Gulland and Holt plot (Pauly, 1984). Arrow or chevron-shaped fish traps offer excellent potential for obtaining fish for tagging in the South Pacific. If traps are well tended, a large number of fish can be obtained over a relatively short period of time for tagging. They can also account for a substantial number of subsequent recaptures, including fish in a condition suitable for multiple release. For details of the construction and operation offish traps see Stott (1970), Munro etal. (1971), Munro (1972), High and Ellis (1973), Munro (1974), Eggers et al. (1982), Sundberg (1985), Ferry and Kohler (1987), Dalzell and Aini (1987), Davies (1989,1992), and Whitelaw etal. (1991). Possibly the most rapidly expanding, and the most contentious, area of literature concerning tropical fisheries science during the last decade relates to estimating growth from analysis of length frequency data. It is also among the oldest, having been developed by Petersen in 1892. The Petersen method assumes that modes evident in large sets of length frequency data are attributable to cohorts of differing age. George and Banerji (1964) developed a variation of this, Modal Progression Analysis, in which the modes in a number of samples accumulated over a known period of time are related, the degree of ad vancement in modes between samples being attributed to growth. Since the development, and subsequent revisions, of the Electronic LEngth Frequency ANalysis (ELEFAN) suite of programs by Pauly and David (1981), a large amount of research effort has been applied both to the development of alternative compu- Shallow Water Reef-associated Finfish 231 ter-based methods of analysis of length data and to applying software that has now been widely distributed, to length data sets. Programs that are now available include those developed by Schnute and Foumier (1980) and Foumier et al. (1990). A comprehensive review and assessment of length-based meth­ ods in fisheries research was conducted during a workshop in Sicily in 1985 (Pauly and Morgan, 1987). Issues associated with the analysis of length frequency data to estimate growth are discussed elsewhere in this volume (see Munro and Moffit). It is noted that length frequency data are relatively easy to collect from South Pacific fisheries although the multispecies and multigear nature of the catch cause difficulties in accumulating representative data sets which constrains subse­ quent analyses. However, Somerton and Kobayashi (1991) have reported on the develop­ ment of new approach for obtaining unbiased estimates of mortality and growth parameters from length-frequency data biased by the size-selectivity of the sampling gear; the Inverse Method for Mortality And Growth Estimation (IMMAGE). Often limitations of application to length data sets have not been recognised resulting in inaccurate assessments of growth parameters. As a general rule, if there is no progression obvious in graphical presentation of the data, it is unlikely that computer analysis will generate realistic results. The most successful applications of computer-based methods to analyse length data have involved small, fast growing, pelagic fishes such as the clupeoids and engraulids (see Dalzell, this volume). Munro and Williams (1985) noted difficulties associated with the compari­ son of growth parameter estimation from different regions and times include the rare incorporation of a procedure for validation, the lack of confidence estimates for derived parameters, the sensitivity of estimates of K to asymptotic size, and the fact that asymptotic size and K are interdependent such that smaller size animals tend to have relatively high values of K. A growth performance index (0 or 0') was developed by Munro and Pauly (1983) and Pauly and Munro (1984) to address this problem. They determined that for species whose growth can be described by the von Bertalanffy equation, it can be demonstrated on empirical and theoretical grounds (Pauly, 1979), that for related animals, double logarithmic plots of the coefficient of growth, K, against the asymptotic weight, W , are linear with a slope of 0.67, such that:

0 = log10 K + 0.67 log10 W„, or

0' =0 log10 K + 2 log|0 LM Thus, at least an initial estimate of K is possible for species for which growth has been reasonably accurately described from other locations. Appeldoorn (1992) asserts that this relationship holds only for intraspecific comparisons and 238 Andrew Wright that for comparisons among species collected using similar methods from the same area, the relationship is more accurately described by: 3> = log K + 0.25 W„

MORTALITY

Estimation of the mortality offish is commonly expressed as: zt N, = N0 e" where N0 and Nt are the numbers offish at time zero and t, respectively and Z is the instantaneous rate of total mortality. Total mortality is the sum of mortality attributable to fishing (F) and natural attrition (M). Thus: Z = F + M Thus when F = 0, Z = M, which means total instantaneous mortality in the absence of fishing is due to natural mortality - the situation that occurs in an unexploited stock (for more detail, see Munro, 1982b; Pauly, 1982; Gulland, 1983). A procedure for estimating the total mortality from the mean size in the catch was suggested by Beverton and Holt (1956). This formulation estimates Z from the mean size of the fish in the catch. Although similar estimates for Z are obtained with this method and that proposed by Ricker (1975) using catch curves, the method of Gulland and Holt is sensitive to changes in mean length. Pauly (1984) discusses some of the constraints associated with this method. Ricker (1975) described the use of the slope of the descending limb on a catch curve, the plot of the natural logarithm offish numbers against their age, to estimate Z. Pauly (1982) described a method for the construction of catch curves from length frequency data. Estimates of natural mortality for tropical fish stocks have generally been obtained from stocks in which no fishing occurs (e.g. Thompson and Munro, 1978; Wright et al. ,1986). For fished stocks, Ricker (1975) proposed a method utilising a time series of Z from the same stock plotted against corresponding values of fishing effort to estimate M as the intercept of the line fitted to these data. This method also provides an estimate of the catchability coefficient (q) based on the slope of the line. Pauly (1980) demonstrated that M can be predicted from a knowledge of growth parameters of a given stock and its mean environmental temperature through an empirical relationship:

loglo M = 0.0066 - 0.279 log10 L^ + 0.6543 Log]0 K + 0.4634 logl0 T where M is the exponential rate of natural mortality, on a yearly basis, L^ is the asymptotic length of the fishi n the stock, in cm, K is their growth coefficient (on a yearly basis) and T (in celsius) is the mean environmental temperature for the stock. According to Pauly (1980), this equation produces reasonable estimates of M for most tropical fish stocks, possibly excluding the small, schooling Shallow Water Reef-associated Finfish 239 pelagics, for which it is slightly over-estimated. However, Pauly (1982) cau­ tioned that estimates of M by this method should be treated as first estimates only as natural mortality of fishes varies with age and is susceptible to predator abundance. Estimates of fishing mortality can be obtained through tag-recapture studies (see Jones, 1977; Ricker, 1975), subtraction of M from Z, for example estimated from length converted catch curves or through Virtual Population Analysis in which the catches in the same year class, or cohort, offish in successive years are considered rather than the numbers caught in different year classes. Jones (1981) has reviewed this and a related method, cohort analysis. Many computer programmes, now incorporate routines for the application of these procedures. Where growth parameters are not available to apply to these procedures, an estimation of the ratio of Z to K, which is closely related to Z (Pauly, 1982), can be estimated from length data alone. This involves estimating hm from the maximum size offish observed in the population. This can be done, with some caution, by assuming that the largest fish sampled is at a size that is 95 per cent of the size of the maximum length attained by that species (Taylor, 1958), or by a procedure known as the Wetherall plot (Wetherall, 1986). With the second procedure, the fish population is assumed to be stable, with constant annual recruitment, von Bertalanffy growth and continuous mortality occurring at a uniform, instantaneous rate. Obviously not all these assumptions can be applied to all fish populations in the tropical Pacific. However, a simple procedure for estimating Z/K can be derived from the equation derived by Beverton and Holt (1956) for estimating Z. This and three other methods, including reservations associated with their application, are described in Pauly (1982). Most procedures to estimate mortality have only been applied to adult fish. Returning to the distinction between recruitment to the habitat from the pelagic environment and recruitment to the fishery, larval survival post- settlement is yet another aspect of reef fish biology for which more information is required. If the rate of early post-settlement mortality is high, and varies in space and time, it is potentially as important as larval mortality while in the plankton in determining the variation measured in rates of recruitment to older age classes (Sale and Ferrell, 1988). In addition, predation on, and competitive interactions among, juveniles may play significant roles in determining the structure of reef fish communities (Smith and Tyler, 1972). Sale and Ferrell (1988) examined this issue for 17 species of pomacentrids, blennids and labrids. For these species, survivorship during the first week following recruitment (= settlement) ranged between 50 and 90 per cent. For eight species mortality was negligible after the first 14 days while in the other 9 species significant mortalities occurred in subsequent weeks. In six species, 75 per cent of individuals survived 45 days while in five others, 20 per cent or 240 Andrew Wright fewer survived that long. Doherty and Sale (1986) reported rates of survival in a reef slope habitat for similar species to those studied by Sale and Ferrell were approximately 65 per cent for the first 30 days of life with most mortality occurring within five days of settlement. Related information for species targeted by island fishermen is not available in the literature.

III. RESOURCE ASSESSMENT

METHODS OF ASSESSMENT

Assessments of fishery harvest from a given area can be divided into three broad groups: those based on a known harvest per unit area, surplus yield assessments and analytical or system models (Munro and Williams, 1985). Marshall (1980a) generalised that coralline shelves with a good cover of live coral, sea grass beds and algae usually can sustain 30-50 kg.ha"1 .yr1 of neritic fishes. Wass (1982) determined that fishing in American Samoa occurred almost exclusively in depths fewer than 8 m, the majority taking place on the reef flat where the annual yield of all marine animals was estimated to be 266 kg.ha" 1 .yr1. The finfish component averaged 106 kg.ha' .yr1 with a range of 69-355 kg.ha"1 .yr1 for different areas fished by villagers within the study site. Russ (1991) noted that yield is a function of fishing intensity and the actual area of reef fished. The issue of the area fished was discussed by Munro and Williams (1985) and Russ (1991) cited examples illustrating the substantial differences in apparent harvests that are generated by considering different areas of operation on reef systems, e.g. to 8 m as in the Samoas, 20-60 m as applied in the 13 studies cited from the Philippines or 30 m as applied in Papua New Guinea. However, in Papua New Guinea, Munro (1976b) used the shelf area to a depth of 200 m to estimate potential yields. Munro and Williams (1985) suggested that the horizontal shelf area between the shelf edge at 20-50 m and the 200 m isobath is negligible although, at least in the South Pacific, no comparisons to substantiate this have been attempted. Differences in the perceived depth of fishing can effectively double the estimated yield (e.g. Bellwood, I988b;Zanne*a/., inprep. cited in Russ, 1991) as can the decision on the degree of affiliation with reefs by the various species components of the fishery, i.e. the demersal, neritic or pelagic-reef transients. Zann et al. {in prep, cited in Russ, ibid.) suggested 40 m as a possible standard as this is the maximum depth of both coral growth and most artisanal fishing techniques in the Pacific. Wright and Richards (1985) selected 30 m on the basis of artisanal fishing practices in Papua New Guinea in combination with the fact that species composition, especially among the lutjanids, lethrinids and serranids, changes significantly around this depth. Shallow Water Reef-associated Finfish 241

The possibility of developing a morpho-edaphic index, developed initially for lakes by Ryder (1965), based on the relative proportions of different habitats in coral reef systems as a basis for predicting theoretical potential yields does not appear to have been investigated further since it was first suggested in 1979 (Roedel and Saila, 1980). Rapid technological developments assisting the interrogation and analysis of aerial photographs and satellite imagery may result in greater use of remote sensing methodologies in resource assessments in future. However, validation by ground surveys will be required for the foresee­ able future to supplement analysis by such techniques (SPC, 1988). Use of surplus production models to estimate potential fish yields has not been extensively applied to reef-associated fisheries in the South Pacific because the time series for catch and effort data are inadequate and production data for a range of fishing effort are generally not available. In addition, surplus production has traditionally been applied to single species fisheries. In the multi-species situation of South Pacific reef fisheries, complex interactions within the community are likely to result in a yield curve for any single species of irregular, not the expected parabolic, shape (Munro and Williams, 1985; Munro, this volume). Recognising this, Munro (1977) and Gulland (1979) used a variant of the Schaefer model to consider the entire catch as a single entity. However, Marten and Polovina (1982) concluded that application of an asymptotic model to data of this sort provided no evidence for any decline in harvest, even at the highest recorded levels of fishing effort. Problems associated with the application of such methods are related to complex issues of catchability and community interactions which remain relatively poorly described for reef fisheries in the South Pacific. The literature contains many examples of the recognition of these problems and various attempts have been made to address them (e.g. Garcia et ai, 1989). A possible alternative to aggregating multi-area or multi-species data to a single production model has been suggested by Polovina (1989). He proposed simultaneously fitting a system of production models to available data using one equation for each area or species. He based this on the assumption that when some parameters can be assumed to be the same for each area or species, different levels of fishing effort among areas could be applied, by simultaneous equations, across the data set available. On the assumption that maximum sustainable yield (MSY) is a function of the unexploited biomass (B J and the turnover rate, which is related to natural mortality (M) in an unfished population, Gulland (1971) proposed: MSY = 0.5 MB. This has been queried by a number of authors (e.g. Caddy and Csirke, 1983). Beddington and Cooke (1983) concluded that this approximation over- 242 Andrew Wright estimates potential yields by a factor which is a function of M. This issue is discussed elsewhere (see Munro, this volume). Application of such estimates to South Pacific fisheries are not common. Somerton and Kobayashi (1990) used a dynamic and equilibrium estimator of spawning potential ratio (SPR) to assess the status of deepwater handline resources in Hawaii. In that location, overfishing is assumed to occur when the spawning stock biomass is 20 per cent of the virgin biomass. Estimates of the virgin biomass of populations of reef fish have been attempted by visual census (Sale, 1980; Sale and Douglas, 1981; Brock, 1982; Sale and Sharp, 1983; Colin et al., 1986; Bohnsack and Bannerot, 1986; Samoilys, 1988. However, the application of visual census methods to species encountered in the catch of island fishermen is difficult due to the extended ranges of many species caught and the cryptic nature of some species (see Doherty et al, 1992). Despite considerable effort expended attempting to develop accurate methodologies for visual assessment of reef community state and composition, the analysis of information obtained using standard parametric procedures has recently been queried (Crimp and Doherty, 1992). Craik (1981a) and Ayling and Ayling (1985) compared the populations of coral trout on fished and unfished reefs on the Great Barrier Reef. The techniques for application of visual assessments of coral trout populations were developed at a series of workshops convened by the Great Barrier Reef Marine Park Authority (Anon. 1985). Bell et al. (1985) and Bellwood and Alcala (1988) reported on the effects offish length on visual estimates offish abundance and biomass and Thresher and Gunn (1986) reviewed the theory and limitations of visual census techniques and reported on six methods of visual census for carangids on the Great Barrier Reef. Unless carefully monitored, it is difficult to assign apparent population changes in such studies solely to the effects of fishing. The possibility of differences between sites resulting from other factors, for example environmental, must also be considered (Russ, 1991). When an assessment of the population within an accurately defined area on a reefprecede s fishing, useful information on standing stock can be attained. Russ and Alcala (1989) reported a significant decrease in the total abundance of many species targeted by local fisheries on a reef in the central Philippines. The families monitored included carangids, caesionids, scombrids, serranids, lutjanids and lethrinids, and it was estimated that lutjanid and lethrinid stocks decreased by 94 per cent after a relatively short period of intensive fishing. Russ (1991) also reported on a study conducted by Beinssen (1988) in which coral trout were tagged in a marine reserve on the Great Barrier Reef prior to fishermen being granted access. Within 14 days of fishing commencing, Beinssen estimated the target species population was reduced by 25 per cent. As noted by Russ {ibid.), this methodology is in effect a depletion experiment, in which total catch, or catch per unit effort, is monitored over a relatively short Shallow Water Reef-associated Finfish 243 period of time. In this period, immigration and emigration are assumed to be negligible, permitting an estimate of virgin standing stock in terms of numbers or weight. The use of sequential catch information over a short period of time to estimate population numbers was developed independently by Leslie and Davis (1939) and DeLury (1947). Polovina (1986) developed this methodology for a multispecies deepwater handline fishery in the Northern Marianas, providing for the catchability for one species to vary inversely with the abundance of competing species. A remaining method for estimating reef fish populations, tagging and recapture, remains to be developed to its potential. This was first attempted by Bardach (1958) and Randall (1962) and has since been rarely applied (but see Parker, 1990; Beinssen, 1988 in Russ, 1991). The theory for estimating popu­ lation size through tag recovery in a single census (Petersen method) or multiple censuses (e.g. the Schumacher and Eschmeyer estimate, Schnabel's method, Ricker's method, or amean of the Petersen method) has been reviewed by Ricker (1975) and Jones (1977) and the assumptions associated with the use of tags have been discussed previously in this chapter. In the South Pacific, the use offish traps offers significant potential for tagging experiments and, if used in conjunc­ tion with another method, for example visual census, could provide relatively accurate assessment of fish population sizes not currently possible using information such as catch and effort which is difficult to collect in this region. Munro and Fakahau (this volume) and Russ (1991) have reviewed appraisal methods and research strategies to assist in fishery and resource monitoring.

YIELDS

A central question to the development of fisheries targeting coral reef- associated organisms concerns the level of fishing finfish communities can sustain. This topic is possibly one of the more complex in the study of reef fisheries. It is dependent on intra- and inter-specific relationships influenced by selective and non-selective removal of components of these extremely diverse communities, the dynamics of which are still not well understood. In addition, among many Pacific islands countries, complex social issues govern marine resource use, thus anthropology and sociology become as important in the study of marine resource exploitation in the region as fisheries or environmental science (e.g. David, 1990). Detailed assessments of the theoretical and actual yields from reef-associ­ ated fisheries commenced in 1974 when Stevenson and Marshall drew attention to high productivity reported for some tropical marine environmental types supporting fisheries (coral reef tracts, 4-10 gm C.nr2.day"'; turtle grass beds, 12 gm Cm2.day"' and mangrove swamps, 12 gm Cm2.day1), and the apparently 244 Andrew Wright high standing stocks offish populations on reefs (380-2,090 kg.ha"1). Atkinson and Grigg (1984) estimated the average net primary production within the diverse habitats of French Frigate Shoals in the Hawaiian chain to be 17 gm Cm" 2.day"'. There are a number of estimates offish standing stocks which confirm ranges between 350 and 1,950 kg. ha'1 (Goldman and Talbot, 1976; Mensavata et al., 1986). Few estimates of the annual production of reef fish communities, as opposed to the annual fisheries harvest, exist. Bardach (1959) estimated annual produc­ tion of a Bermudan patch reef community at 176 kg. ha"1 which was approxi­ mately 35 per cent of the standing crop. On this basis, Montgomery (1990) deduced that reefs supporting a standing stock of 2,000 kg. ha"1 should produce 700 kg.ha"1 .yr1. Actual fisheries harvests from coral reefs have been reviewed by Marshall (1980a, 1980b), Marten and Polovina (1982), Russ (1984,1991) and Munro and Williams (1985). Munro and Williams (1985, Table I) and Russ (1991, Table I) collated available information on fisheries harvests from reef areas in the Indian Ocean, Caribbean and the Pacific. Estimates for the total annual harvest of marine resources in the South Pacific region have been attempted in American Samoa (Hill, 1977; Wass, 1982), Western Samoa (Zann et al, in prep, in Russ, ibid.), Kiribati (Mariott, 1984) and Papua New Guinea (Wright and Richards, 1985; Dalzell and Wright, 1986; Lock, 1986). These have ranged from less than 5 kg.ha"1 .yr'1. in northern Papua New Guinea (Wright and Richards, 1985) to 50 kg.ha"1 .yr1. close to the national capital of Papua New Guinea (Lock, 1986), to more than 110 kg.ha1.yr'. in Western Samoa (Zann et al, in prep, in Russ, 1991). Mariott (1984) reported a yield of about 44 kg.ha"' .yr"1 from the reefs and lagoon in Tarawa Lagoon, Kiribati. Alcala (1981) and Alcala and Luchavez (1981) reported yields of 80-237 kg.ha'.yr' for reefs extending to depths of 60 m surrounding Apo and Sumilon Islands in the Philippines. At Apo Island, also in the Philippines, Bellwood (1988b) reported the 39 families offish harvested, produced an annual estimated yield of 248.6 kg.ha"'. A trophodynamic model for fish biomass production on a reef tract on the Great Barrier Reef suggests that the potential fish production relative to primary production from the reef studied is as high as for other productive marine ecosystems (Polunin and Klumpp, 1991). At Davies Reef, these researchers used the model to estimate the potential fishery yield for macro-grazers over the whole reef to be 40 kg.km'2, invertebrate-feeders 70-160 kg.knr2, piscivores 0- 20 kg.km2 and planktivores 30 kg.km2 with the maximum predicted yield for the reef estimated to be 140-230 kg.km2 for a fishery targeting macro-grazing, invertebrate-feeding and planktivorous fishes. Lock (1986) reported that a relatively high yield on the Port Moresby reefs was due to a combination of the proximity of the reef fishery to a densely populated urban centre and the fact that the fishermen were residents of low Shallow Water Reef-associated Finfish 245 islands off the mainland with few alternative means to generate cash than fishing. A similar situation applies at the Kiribati site. Wright and Richards (1985) suggested that the relatively low yield of finfish from the reefs in the Tigak Islands in northern Papua New Guinea was likely to be indicative of most reef fisheries in Papua New Guinea. The reason for this was ascribed to the fact that most mainland villagers with access to productive agricultural land prefer to garden. Fishing is considered a leisure-time activity in these situations (Wright and Kurtama, 1988; Leqata et al. 1990; Dalzell and Wright, in press). However, on low islands and atolls in Papua New Guinea and elsewhere in the region, fishing is the dominant activity.

IV. FISHERIES DESCRIPTION

A number of factors affect the ability of biologists to monitor coral reef fisheries in the Pacific islands region. These include the dispersed nature of fishing operations and landing sites, the use of a variety of gears during fishing expeditions and the associated difficulty of assigning portions of the catch by gear type, taxonomic difficulties with respect to accurate identification of an extremely diverse catch, the intermittent involvement of many participants, and difficulty in measuring effort (for example see Stevenson and Marshall, 1974; Wright and Richards, 1985; Bellwood, 1988b; Russ, 1991). In addition, fish poisoning or ciguatera poses particular problems for researchers in the South Pacific. In many areas of Micronesia and Polynesia in particular, the presence of ciguatoxic fish is a major factor influencing fishing activity in terms of both locations fished and the composition of the catch retained and returned to the landing site (e.g. Hiyama and Yasuda, 1975; Anderson and Lobel, 1987; Randall, 1980; Bagnis, 1992). In order to gain realistic understanding of reef-associated fisheries in the Pacific, it is important to monitor subsistence fisheries in conjunction with the artisanal fishery. In many situations the catch monitored from the artisanal fishery is confined to that portion of the catch presented for sale. Often a portion of the catch, mostly consisting of smaller fishes, is retained for home consump­ tion thus is not presented at a market point therefore is often not recorded in catch sampling exercises. Many Pacific island countries rank among the highest in the world for the per capita consumption offish with some atoll nations consuming up to 250 kg per year. Even the relatively urbanised and agricultural islandssuch as Fiji consume 45 kg per head annually. In addition, fishing effort and gears vary between artisanal and subsistence fishing activities, with nets for example, being only employed to provide subsistence needs on the occasion of a celebration in the village. The dominance of handlines and spearfishing in the subsistence catch, which is mainly taken 246 Andrew Wright from inshore waters, results in a significantly different catch composition when compared with artisanal catches dominated by net and troll gear. In the Tigak Islands in northern Papua New Guinea, Wright and Richards (1985) noted that 20 per cent of the subsistence catch was taken by spear and a similar proportion of the catch presented for sale at the municipal market was assumed to have been also taken by this method. In Solomon Islands where a fishing competition was organised to gain access to village catch data, it was estimated that spearing accounted for 14 per cent of the catch and that this was representative of the catch by this method in both artisanal and subsistence fisheries (Blaber et al. 1990a; b). In American Samoa, Hill (1977) and Wass (1982) respectively estimated that spearing accounted for 26.4 and 31.0 per cent of the total catch monitored. A similar situation exists with analysis of the handline catch from the Tigak Islands. In the artisanal fishery, handlines accounted for 21.6 per cent of the catch, a similar proportion as monitored in the fishery in American Samoa. The majority of the subsistence catch in the Tigak Islands is also handlined, thus it was estimated that this method accounted for 42 per cent of the total catch (subsistence and artisanal fisheries) in these islands (Wright and Richards, 1985). The percentage of the catch estimated as handlined in Roviana and Tulagi Lagoons in Solomon Islands was 61.5 (Blaber et al. 1990c). Wright and Richards (1985) noted that traditional fishing craft in the Tigak Islands do not provide a stable fishing platform in unsheltered waters, and for this reason, together with the relatively high cost of fuel, trolling was not recorded as a common method of fishing. However, this is not the case for all island groups in the South Pacific and in Solomon Islands, for example, Blaber et al. (1990c) reported trolling accounted for approximately 23 per cent of subsistence and artisanal catches. Trolling with artificial lures or fish baits is widely practiced throughout the region with the catch being dominated by large scombrids such as the Spanish mackerel, Scomberomorus commerson, sphyraenids, thunnids and carangids. During a 13-month investigation of subsistence and artisanal fisheries in the Tigak Islands of northern Papua New Guinea, Wright and Richards (1985) recorded 253 species from the catch sampled. These species represented fish with habitats ranging from strongly reef-associated demersal habitats to neritic pelagic and near-shore oceanic. Hiatt and Strasburg (1951) recorded 175 species of fishes and invertebrates in the catches of the residents of Arno Atoll in the Marshall Islands despite no fishing for pelagic species. Banner and Randall (1952) recorded 396 species of fishes taken by residents of Onotoa in Kiribati, a large portion of which were consumed. The limited periods of these studies did not permit an estimate of total fish harvest, but on both atolls, the annual harvest was concluded to be significant. Blaber et al. (1990c) recorded 183 species from 31 families in the catch landed for five fishing competitions in Shallow Water Reef-associated Finflsh 247

Solomon Islands. However, early in the 1990s it appears that specialised fisheries targeting a limited number of species has potential to develop rapidly in the region. These fisheries, targeting serranids and the Maori wrasse, Cheilinus undulatus are producing live fish sea-freighted to Asian markets. The first operation commenced in Palau in the late 1980s and by 1992 fisheries had extended through Papua New Guinea as far east as Tuvalu. Demersal, reef-attached species dominated the catch monitored in the Tigak Islands in northern Papua New Guinea (Wright and Richards, 1985) and the lagoon fishery in New Caledonia (Loubens, 1978), contributing 83.5 per cent and 80 per cent respectively to the recorded catches. From data provided by Wass (1982) in American Samoa, demersal fishes contributed 68 per cent to the nearshore catch there. The species composition in the Tigak Islands differed somewhat to that recorded from American Samoa. This is not unexpected as the fish fauna of American Samoa is not as diverse as in Papua New Guinea, which is close to the area of greatest species diversity in the Philippine and Indonesian Archipelagos. In addition, American Samoa's main island is high, whereas the PapuaNew Guinea study site concentrated in a shallow lagoon surrounded by many low islands. Although Hill (1977) recorded lethrinids from the catches monitored in American Samoa, Wass (1982) noted their absence. In the Tigak Islands, 17 species of lethrinids contributed 10 per cent by weight to the artisanal catch (Wright and Richards, 1985). Grant (1981) reported 5-40 per cent of the dropline catch on the Great Barrier Reef were lethrinids, and Loubens (1978) reported this family contributed 39 per cent to the New Caledonian lagoon fishery. Lethrinids were the dominant demersal species monitored at municipal markets in Fiji during 1990, contributing 15 per cent to landings which were dominated by pelagic species such as scombrids and sphyraenids (Anon. 1991). Although monitored landings of serranids were depressed rela­ tive to previous years, the composition of the catch landed at municipal markets in Fiji during 1990 was similar to previous years, approximately 10 per cent. From data presented by Wass (1982), carangids (principally the scad, Selar crumenophthalmus), dominated the American Samoan catch, accounting for 32 per cent of recorded landings. Acanthurids (19 per cent), serranids (11 per cent), lutjanids (7 per cent) and scarids (5 per cent) comprised the other major groups. Bellwood (1988b) reported the fish catch from Apo Island in the Philippines was composed of 38 families. He noted the catch was dominated by relatively few families, with nine families contributing over 93 per cent of the total annual yield, 40 per cent of which was contributed by carangids and 23.6 per cent by acanthurids. He noted little variation throughout the year with respect to the dominance of acanthurids and carangids in the catch, although within season variability among most families was evident. Bellwood noted the major differ­ ence between the catch recorded at Apo Island and that recorded at Sumilon 248 Andrew Wright

Island by Alcala (1981) was the dominance of caesionids in the Sumilon catch. Bellwood (1988b) also noted the limited value of familial composition in determining the nature of the fish yield and factors which determine its size. He suggested a clearer understanding of reef-associated fisheries could be achieved if fish are grouped by their degree of affiliation with the reef itself. He noted acanthurids, for example, include reef-feeding and planktivorous species. Similarly, the lutjanids, scarids, carangids, lethrinids, siganids and balistids include species that range from distinctly reef-resident to neritic pelagic. At Apo Island, Bellwood (1988b) recorded the dominant proportion of the fish catch (56.6 per cent) was not dependent on the reef for food or shelter. Russ (1991) observed that many fisheries on coral reefs target fishes lower in the food chain. He cited examples from French Polynesia (Stein, 1988), Palau (Johannes, 1981 a), PapuaNew Guinea (Wright and Richards, 1985; Dalzell and Wright, 1986) and the Cook Islands. Parrot fish have probably been taken by residents of Palmerston Atoll in the Cook Islands since it was first settled in 1860. For the majority of this period, the fish were taken by coconut frond scare lines (Sims, 1988; Anon., 1989). Since 1985 a fishery consisting predomi­ nantly of four species common on reef flats has operated using nylon nets to block escape channels from the reef flat to the reef face on rising or falling tides. Sims (1988) estimated the annual harvest of parrotfish from this fishery to be 15-20 tonnes. In other cases, such as reported from the central Philippines, catches ofplanktivores are high (e.g. Alcala, 1981; AlcalaandLuchavez, 1981; Bellwood, 1988b). The breakdown of the catch recorded from the Tigak Islands (Wright and Richards, 1985) by broad trophic grouping is presented in Table III.

V. MANAGEMENT CONCERNS

It has been noted elsewhere (SPC, 1988; Munro and Fakahau, this volume), that there have been few attempts to manage marine resource exploitation in the South Pacific. Although some fisheries for beche-de-mer are traditionally managed, e.g. the fishery operating at Ontong Java in Solomon Islands, and quota systems have been applied to some trochus fisheries in the Marshall Islands, the Federated States of Micronesia and the Cook Islands, there are few contemporary examples of management of fisheries for finfish in the South Pacific region in the literature. However, island communities have practiced traditional forms of management, through limited entry applied under reef, lagoon and marine resource tenure systems, probably since the settlement of the island groups. It has only been relatively recently that established traditional systems have been considered as possible mechanisms to address present-day management issues (Johannes, 1981b; Hviding, 1990). Although few attempts at this have been documented, Sims (1990) describes application of established communal practices to manage a trochus fishery at Shallow Water Reef-associated Finfish 249

Aitutaki in the Cook Islands and Wright (1990) and Turner (1991) discuss various issues associated with the distribution of bait royalty payments in a fishery for live bait for pole-and-line tuna fishing in northern Papua New Guinea. Others note that the abundances and average sizes of many large carnivores increase within protected areas and that smaller fishes and species with different trophic levels show similar patterns where they are the focus of fishing but that there remain a number of key questions regarding the function of reserves in fishery management, and as a result, the ability of traditionally- based systems to provide effective conservation mechanisms in the face of Table III. The composition of the small-scale reef-associated fishery catch in the Tigak Islands, Papua New Guinea by trophic group (after Wright and Richards, 1985).

Piscivores Omnivores Herbivores 30.0% 31.8% 38.2%

Serranids(3.0%) Lethrinids(10.4%) Mugilids(21.2%) Sphyraenids(

VI. CONCLUSION

Environmental, resource, economic and social issues all require detailed consideration in the management of the exploitation of reef resources, thus the need to involve expertise capable of providing assessments of a multidisciplinary nature requires greater consideration in South Pacific fishery resource manage- 252 Andrew Wright ment initiatives. At present, although work occurs among these individual disciplines in the region, much remains to be done to promote a coordinated, multidisciplinary approach. As is evident from the attached list of references, the literature dealing with the biology, ecology and fisheries for reef-associated fishes is substantial and now rapidly expanding. This chapter has attempted to introduce much of the literature specific to South Pacific fisheries. However, some issues are only briefly mentioned. If expansion on the foregoing is desired, researchers and students alike are referred to three recent comprehensive edited reviews that consolidate most recent information for marine fish biology and population ecology: one concerned with reef fish trophic and larval ecology, reproductive patterns and community organisation, including a review of reef-associated fisheries incorporating issues requiring further research (Sale, 1991), a second covering zoogeography, behavior and ecology (Montogomery, 1990), and last a book describing methods in fisheries research (Schreck and Moyle, 1990).

NOTE 1. Moffit,R.B.u/i/>«W. manuscript. Age, growth and reproduction of the k\m\u,Parupeneus porphyreus (Jenkins). Directed Research Report, Master of Science, Plan B. Depart­ ment of Zoology, University of Hawaii. September 1979.

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