217
Capture-based aquaculture of
groupers
Mark Tupper
WorldFish Center
Penang, Malaysia
E-mail: [email protected]
Natasja Sheriff
WorldFish Center
Penang, Malaysia
E-mail: [email protected]
Tupper, M.; Sheriff, N. 2008. Capture-based aquaculture of groupers. In A. Lovatelli
and P.F. Holthus (eds). Capture-based aquaculture. Global overview. FAO Fisheries
Technical Paper. No.  508. Rome, FAO. pp. 217–253.
SUMMARY
The economies of China and Southeast Asia have developed rapidly over the past two
decades, leading to the emergence of a wealthy class with substantial disposable income.
This has led to an increasing demand for fish in the region (Birkeland, 1997). The “live
fish trade” of the Indo-Pacific has expanded rapidly in recent years, and now targets many
species (Johannes and Riepen, 1995; Sluka, 1997, Sadovy and Vincent, 2002). Groupers
are greatly valued for the quality of their flesh, and most species command high market
prices. Groupers are the most intensively exploited group in the live fish trade, and the
high prices paid by exporters to local fishermen mean that target species may be heavily
over-fished (Morris, Roberts and Hawkins, 2000). In order to alleviate the pressure on
wild grouper stocks, many nations have promoted aquaculture in the hopes of producing a
more sustainable grouper yield. However, full-cycle culture of most grouper species is not
yet possible, although several important advances have been made in recent years. For this
reason, about two-thirds of all grouper culture involves the capture and grow-out of wild
seed (Sadovy, 2000). This is known as capture-based aquaculture (CBA).
There are at least 16 species of groupers that are cultured in many Southeast Asian
countries, including Indonesia, Malaysia, Philippines, Taiwan Province of China,
Thailand, China Hong Kong Special Administrative Region (SAR), the southeast of the
China and Viet Nam (Sadovy, 2000). Grouper culture is also undertaken in India, Sri
Lanka, Saudi Arabia, Republic of Korea, Australia, the Caribbean and in the southeastern
United States of America. Despite the huge popularity of live fish in China and Southeast
Asia, only 15–20  percent of the amount consumed each year comes from aquaculture, as
culture is principally constrained by limited and unreliable supplies of wild seed and the
difficulties of spawning in captivity.
Grouper seed is collected using a variety of methods. Capture methods are generally
artisanal and the fishermen employ a variety of artificial habitats. Some grouper seed
collection methods are more damaging than others. Clearly destructive methods include
those that result in high mortality, involve high levels of bycatch, and/or cause damage
to the fish habitat. A further problem is that some methods result in monopolization of

Capture-based aquaculture: global overview218
the local fishery by a few individuals. Destructive methods include scissor nets and fyke
nets, which are already banned in some areas. The mortality rates that follow capture and
transport are not well documented; estimates for over the first 2 months after harvest
are quite variable (30–70  percent), depending on the quality of fry, the level of transport
stress, and the presence of disease and cannibalism (Pudadera, Hamid and Yusof, 2002).
Because full-cycle culture of most grouper species is not yet possible, approximately
66–80  percent of all grouper culture involves the capture and grow-out of wild seed and
the volume of seed caught each year exceeds hundreds of millions of individuals (Sadovy,
2000). When seed catches are compared to the numbers of marketable fish produced, the
results strongly suggest crude and wasteful culture practices. Sadovy (2000) estimated
that about 60 million seed fish are needed to produce the regional total of 23  000 tonnes
of table-size live fish from culture annually.
Trash fish is commonly used for feeding in grouper cage culture, but its increasing
cost, shortage of supply, variable quality and poor feed conversion ratios indicate that this
form of feed may not be the best from either a nutritional or an economic point of view.
A dependable supply of cost-effective, non-marine, sources of alternative protein must
be provided if grouper farming is to remain profitable. Millemena (2002) demonstrated
that up to 80  percent of fishmeal protein can be replaced by processed meat meal and
blood meal derived from terrestrial animals with no adverse effects on growth, survival,
and food conversion ratio (FCR). From an economic standpoint, replacement of fishmeal
with cheaper animal by-product meals in practical diets can alleviate the problem of low
fishmeal availability and high costs.
Recent research suggests that the ecological footprint of capture-based grouper
aquaculture is large (Mous et al., 2006). Support for grouper CBA is often based on the
assumption that the natural morality of early juvenile grouper is very high, so that the
fishery is not adding substantially to this natural mortality and therefore not affecting
adult population size to any great extent. This assumption remains untested for most
grouper species. However, recent research suggests that the period of very high mortality
occurs during and immediately after settlement, and that juvenile grouper surviving more
than a few days have a much higher chance of survival (Tupper, 2007). In addition to
problems of bycatch, wasteful mortality, and overfishing, cage and net culture can create
other environmental problems, most notably point-source pollution which can have
adverse effects on coastal waters, and particularly on coral reefs.
As a contributor to rural livelihoods, particularly those of coastal fishers, grouper
aquaculture can generate potentially large financial benefits. The high value of grouper
on the export market ensures that farmers are able to generate a profit even when stocks
suffer heavy mortalities. Despite high initial investment costs, studies have shown that
with appropriate support, even the poorest can benefit from grouper culture, with
implications for both household well-being and community development. However,
based on the information reviewed in this report, capture-based aquaculture may not be
the best means to ensure a steady and sustainable supply of grouper for either the live or
“non-live” fish trades. This is due to a number of problems including low availability of
seed, destructive and wasteful seed collection techniques, removal of large numbers of
early life history stages with subsequent impacts on adult populations and conflicts with
capture fisheries, and pollution and disease resulting from culture operations.
The obvious solution to some of the problems of CBA for grouper is to develop
closed-cycle hatchery rearing for all the grouper species sought by the market. Important
advances in full-cycle culture have been made for several species, particularly in
Taiwan  Province of China, and full-cycle culture appears financially feasible given a large
enough capital investment. However, given the financial means of most grouper culturists,
and the difficulty in rearing most grouper species, it remains unlikely that many of these
species will be hatchery-reared in the near future. In the meantime, steps must be taken
to improve the management of both CBA and capture fisheries for grouper.

Capture-based aquaculture of groupers 219
INTRODUCTION
The economies of China and Southeast Asia have developed rapidly over the past two
decades, leading to the emergence of a wealthy class with substantial disposable income.
This has led to an increasing demand for fish in the region (Birkeland, 1997). The “live
fish trade” of the Indo-Pacific has expanded rapidly in recent years, and now targets
many species (Johannes and Riepen, 1995; Sluka, 1997; Sadovy and Vincent, 2002).
Groupers are greatly valued for the quality of their flesh, and most species command
high market prices. Groupers are the most intensively exploited group in the live fish
trade, and the high prices paid by exporters to local fishermen mean that target species
may be heavily over-fished (Morris, Roberts and Hawkins, 2000). Trade often follows
a pattern of sequential over-exploitation; the most highly sought species are fished-
out in country after country, before the less valuable species are targeted and fished
intensively (Sluka, 1997; Johannes and Riepen, 1995). Wealthy customers pay very
high prices for endangered species in Chinese and Southeast Asian markets. In 1997
the red grouper, Epinephelus akaara, fetched US$42/kg in China Hong Kong SAR
markets. In 2004, restaurants were charging US$225 for only the lips of the humphead
wrasse, Cheilinus undulatus. Thus, fishermen will go to great lengths in order to catch
every fish, and this has already contributed to regional population crashes of species,
including Epinephelus akaara and Epinephelus striatus (Morris, Roberts and Hawkins,
2000; Sadovy, 2001a).
The impact of intensive fishing is exacerbated by the K-selected life strategies of
these genera, their tendency to form predictable spawning aggregations and their
occurrence on relatively shallow, easily accessible coral reefs, which are severely over-
exploited in many parts of the world. For many of these species, spawning aggregations
represent the total reproductive output for a given year, and many species consistently
return to the same aggregation area, year after year. Fisheries often target spawning
aggregations, since they are consistent in time and space and large numbers of fish can
easily be caught in a short time (Rhodes and Tupper, 2007). When fishing pressure
removes a high proportion of the fish forming these aggregations, these may quickly
decline, and within a few years may cease to form altogether (Johannes et al., 1999;
Sadovy and Eklund, 1999).
A large proportion of the world’s groupers are caught in artisanal fisheries, and
even low-level artisanal fisheries can adversely affect stocks of these highly vulnerable
species. Recreational fishing may also have significant impact on stocks; for example,
the recreational fishery of groupers accounts for up to 35  percent of Florida’s (United
States of America) total grouper catch (Morris, Roberts and Hawkins, 2000). The
global catch of groupers showed a 68  percent increase from 100  724 tonnes in 1991 to
168 943 in 2000. In order to alleviate the pressure on wild grouper stocks, many nations
have promoted aquaculture in the hopes of producing a more sustainable grouper yield.
Because grouper are particularly difficult to culture in closed systems, full-cycle culture
of most grouper species is not yet possible (although several important advances have
been made in recent years). For this reason, about two-thirds of all grouper culture
involves the capture and grow-out of wild seed (Sadovy, 2000). This is known as
capture-based aquaculture (CBA).
There is a strong link between fishing activity and the capture-based seed used
for farming, with declines in premium species from the overfishing of grouper
adults. However, the reasons for this decline cannot be evaluated without careful,
controlled studies, as falling catches may in fact be due to a combination of different
causes: overfishing of the adults which produce the juveniles, habitat degradation and
pollution, destructive fishing techniques, high export demand, etc. (Johannes, 1997;
Sadovy, 2000). A more holistic management approach to establish the links between
adults and juveniles is necessary.

Capture-based aquaculture: global overview220
SPECIES DESCRIPTIONS AND THEIR USE IN AQUACULTURE
Groupers (class Actinopterygii, order Perciformes, family Serranidae, sub-family
Epinephelinae) comprise 14 genera and 449 species of the subfamily Epinephelinae, or
roughly half of all species in the family Serranidae (groupers and sea basses) (Heemstra
and Randall, 1993). There are 16 major grouper species that are cultured; the dominant
species vary somewhat regionally. The most consistently abundant species that are
captured for culture purposes and also reared in hatcheries are Epinephelus coioides
and E. malabaricus. Other important species are E. bleekeri, E. akaara, E. awoara
and E. areolatus. E. amblycephalus, E. fuscoguttatus, E. lanceolatus, E. sexfasciatus, E.
trimaculatus, E. quoyanus, E. bruneus, Cromileptes altivelis, Plectropomus leopardus
and P. maculatus are cultured in small amounts. In the southeastern United States
of America and the Caribbean, E. striatus, E. itajara, Mycteroperca microlepis and
M. bonaci seem to have good farming potential (Tucker, 1999). However, CBA for
groupers in the western hemisphere has not been developed to any large extent, unlike
in Southeast Asia.
Juveniles and adults of some grouper species live in coastal or lagoonal waters and
estuaries, while others prefer the cleaner waters of offshore reefs. Their eggs are single,
non-adhesive, and buoyant at normal salinities. The larvae of most species spend
about 30–50 days as planktonic larvae (Colin, Koenig and Laroche, 1996). As they
become juveniles, groupers settle in shallow waters where they seek shelter in seagrass
beds, mangrove prop roots, coral rubble, branching coral or branching macroalgae.
Some juvenile groupers are habitat generalists, settling in any available shelter, while
other species have specific nursery habitats in which their growth and survival are
enhanced (Tupper, 2007). After hatching, wild grouper larvae eat copepods and other
small zooplankton. They switch to larger crustaceans, such as amphipods and mysid
shrimp, as they grow. Wild juveniles and adults eat fish, crabs, shrimp, lobsters and
molluscs (Tucker, 1999), although the genus Plectropomus tends to be predominantly
piscivorous.
Groupers range in maximum size from only 12 cm (e.g. Paranthias colonus) to over
3 m (e.g. Epinephelus lanceolatus). Most groupers that have been studied are sexually
mature within 2–6 years, but some of the larger species may take longer to mature,
e.g. Epinephelus fuscoguttatus, which matures at about 9 years. Most serranids are
protogynous hermaphrodites. As a rule, some change from female to male as they grow
older; others may change only if there is a shortage of males. In nature, many species
spawn in large aggregations (hundreds to thousands of fish) with a sex ratio nearing
1:1 (Rhodes and Sadovy, 2002). In some cases, several grouper species may share the
same aggregation site (e.g. in Palau and Pohnpei; see Johannes et al., 1999; Rhodes and
Tupper, 2007).
Groupers are some of the top predators on coral reefs, and tend to be K-strategists
demonstrating slow growth, late reproduction, large size and long life-spans which
make them vulnerable to overexploitation. Also contributing to their vulnerability
is the fact that they are sex-changers with a low proportion of males in the smaller
cohorts, which means that heavy fishing pressure often removes most of the males
(or removes fish before they can become male). Additionally, many groupers form
spawning aggregations that are predictable in space and time, making them extremely
easy to harvest. These aggregations can represent the entire annual reproductive output
for some species. Groupers are sedentary in character and strongly territorial, making
them easy targets for spear fisheries (Bullock et al., 1992; Heemstra and Randall, 1993;
Sadovy, 1996; Domeier and Colin, 1997; Sadovy and Eklund, 1999; Morris, Roberts
and Hawkins, 2000). Tables 1–16 summarize the characteristics of grouper species most
commonly encountered in CBA, while Figures 1–32 illustrate their appearance and
geographical distribution.

Capture-based aquaculture of groupers 221
FIGURE 2
Distribution of Cromileptes altivelis (FishBase, 2007)
TABLE 1
Characteristics of the humpback grouper, Cromileptes altivelis
Common names: Humpback grouper, panther grouper, mouse grouper, highfin grouper
Size and age: Max size 70.0 cm TL
Environment: Reef-associated; marine; depth range 2–40 m
Climate: Tropical; 32°N - 23°S, 88°E - 168°E
Importance: Juveniles are commonly caught for the aquarium trade while adults are utilized as a food
fish. Very high value in China Hong Kong SAR live fish markets.
Resilience: Low, minimum population doubling time 4.5–14 years.
Biology and ecology: Generally inhabits lagoon and seaward reefs and are typically found in dead or silty areas.
Also found around coral reefs and in tide pools. Growth is very slow. Feed on small fishes
and crustaceans.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF J.E. Randall
FIGURE 1
Humpback grouper (Cromileptes altivelis)
Cromileptes altivelis (Valenciennes, 1828)

Capture-based aquaculture: global overview222
FIGURE 4
Distribution of Epinephelus akaara (FishBase, 2007)
TABLE 2
Characteristics of the Hong Kong grouper, Epinephelus akaara
Common names: Hong Kong grouper
Size and age: 53.0 cm TL; max. published weight: 2  470 g
Environment: Reef-associated; marine
Climate: Tropical; 39°N - 20°N, 109°E - 143°E
Importance: A highly prized food fish in China Hong Kong SAR live fish markets.
Resilience: Medium, minimum population doubling time 1.4–4.4 years.
Biology and ecology: Little is known about the biology and ecology of this species. Usually caught by hand-lining
over rock strata. Listed as endangered by IUCN Grouper And Wrasse Specialist Group.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF A. Cornish
FIGURE 3
Hong Kong grouper (Epinephelus akaara)
Epinephelus akaara (Temminck and Schlegel, 1842)

Capture-based aquaculture of groupers 223
FIGURE 6
Distribution of Epinephelus amblycephalus (FishBase, 2007)
TABLE 3
Characteristics of the banded grouper, Epinephelus amblycephalus
Common names:Banded grouper
Size and age: 50.0 cm TL
Environment: Reef-associated; marine; depth range 80–130 m
Climate: Tropical; 35°N - 20°S, 95°E - 179°W
Importance: Fisheries: minor commercial.
Resilience: Medium, minimum population doubling time 1.4–4.4 years.
Biology and ecology: Little known.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF CSIRO
FIGURE 5
Banded grouper (Epinephelus amblycephalus)
Epinephelus amblycephalus (Bleeker 1857)

Capture-based aquaculture: global overview224
FIGURE 8
Distribution of of Epinephelus areolatus (FishBase, 2007)
TABLE 4
Characteristics of the areolate grouper, Epinephelus areolatus
Common names: Areolate grouper
Size and age: 47.0 cm TL; max. published weight: 1  400 g; max. reported age: 15 years
Environment: Reef-associated; marine; depth range 6–200 m
Climate: Tropical; 35°N - 33°S, 29°E - 180°E
Importance: An important fisheries and aquaculture species in the Live Reef Fish Trade (LRFT).
Resilience: Medium, minimum population doubling time 1.4–4.4 years.
Biology and ecology: Usually found in seagrass beds or on fine sediment bottoms near rocky reefs, dead coral,
or alcyonarians, in shallow continental shelf waters. Juveniles are common at water depths
to 80 m. Probably spawn during restricted periods and form aggregations when doing
so. Eggs and early larvae are probably pelagic. Feed on fish and benthic invertebrates,
primarily prawns and crabs.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF J.E. Randall
FIGURE 7
Areolate grouper (Epinephelus areolatus)
Epinephelus areolatus (Forsskål, 1775)

Capture-based aquaculture of groupers 225
FIGURE 10
Distribution of Epinephelus awoara (FishBase, 2007)
TABLE 5
Characteristics of the yellow grouper, Epinephelus awoara
Common names: Yellow grouper
Size and age: 60.0 cm TL
Environment: Reef-associated; marine; depth range 10–50 m
Climate: tropical; 39°N - 12°N, 110°E - 143°E
Importance: Commercial fisheries and aquaculture; medium value in China Hong Kong SAR live fish
markets.
Resilience: High, minimum population doubling time less than 15 months (Fecundity  = 24 329).
Biology and ecology: Occurs in rocky areas as well as on sandy-mud bottoms. Juveniles are common in tide pools.
In captivity, the species is aggressive, chasing and biting other species, especially members
of its own species. Protogynous hermaphrodite. Artificial fertilization of eggs was done
and the longest survival time for the larvae was 15 days.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF D.C. Cook
FIGURE 9
Yellow grouper (Epinephelus awoara)
Epinephelus awoara (Temminck & Sclegel 1842)

Capture-based aquaculture: global overview226
FIGURE 12
Distribution of Epinephelus bleekeri (FishBase, 2007)
TABLE 6
Characteristics of the duskytail grouper, Epinephelus bleekeri
Common names: Duskytail grouper
Size and age: 76.0 cm TL
Environment: Demersal; marine; depth range 30–104 m
Climate: Tropical; 32°N - 17°S, 48°E - 136°E
Importance: Minor commercial fisheries value, moderate commercial aquaculture value. In China Hong
Kong SAR live fish markets.
Resilience: Low, minimum population doubling time 4.5–14 years (t max=24).
Biology and ecology: Occurs on shallow banks, but is not known from well-developed coral reefs. Usually taken
by trawling in 30–45 m or by hand-lining over rocky banks.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF J.E. Randall
Epinephelus bleekeri (Temminck & Sclegel 1842)
FIGURE 11
Duskytail grouper (Epinephelus bleekeri)

Capture-based aquaculture of groupers 227
FIGURE 14
Distribution of Epinephelus bruneus (FishBase, 2007)
TABLE 7
Characteristics of the longtooth grouper, Epinephelus bruneus
Common names: Longtooth grouper
Size and age: 128 cm TL (male/unsexed; Ref. 40637); max. published weight: 33.0 kg (Ref. 40637)
Environment: Reef-associated; marine; depth range 20–200 m
Climate: Tropical; 38°N - 17°N, 108°E - 142°E
Importance: Important in commercial and recreational fisheries. Commercially cultured in Japan and
China Hong Kong SAR.
Resilience: Very low, minimum population doubling time more than 14 years.
Biology and ecology: Inhabits rocky reefs; also found on muddy grounds. Juveniles occur in shallow waters.
Source: Modified from FishBase (Froese and Pauly, 2007).
FAO
Epinephelus bruneus (Bloch, 1793)
FIGURE 13
Longtooth grouper (Epinephelus bruneus)

Capture-based aquaculture: global overview228
FIGURE 16
Distribution of Epinephelus coioides (FishBase, 2007)
TABLE 8
Characteristics of the orange-spotted grouper, Epinephelus coioides
Common names: Orange-spotted grouper, estuary grouper, green grouper
Size and age: 120 cm TL (male/unsexed; Ref. 47613); max. published weight: 15.0 kg (Ref. 11228); max.
reported age: 22 years
Environment: Reef-associated; brackish; marine; depth range 2–100 m
Climate: Subtropical; 37°N - 34°S, 28°E - 180°E
Importance: Important for commercial fisheries and aquaculture throughout Southeast Asia; major
species in China Hong Kong SAR live fish markets.
Resilience: Medium, minimum population doubling time 1.4–4.4 years (K=0.17; tmax=22).
Biology and ecology: Inhabit turbid coastal reefs and are often found in brackish water over mud and rubble.
Juveniles are common in shallow waters of estuaries over sand, mud and gravel and among
mangroves. Feed on small fishes, shrimps, and crabs. Probably spawn during restricted
periods and form aggregations when doing so. Eggs and early larvae are probably pelagic.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF D.C. Cook
Epinephelus coioides (Hamilton, 1822)
FIGURE 15
Orange-spotted grouper (Epinephelus coioides)

Capture-based aquaculture of groupers 229
FIGURE 18
Distribution of Epinephelus fuscoguttatus (FishBase, 2007)
TABLE 9
Characteristics of the brown-marbled grouper, Epinephelus fuscoguttatus
Common names: Brown-marbled grouper, tiger grouper, dusky grouper, flowery grouper, flowery cod
Size and age:  120 cm TL; max weight 35.0 kg, max. age >40 years
Environment: Reef-associated; marine; depth range 1–60 m
Climate: Tropical; 35°N - 27°S, 39°E - 171°W
Importance: Minor commercial fisheries, moderate importance in aquaculture and live reef fish trade.
Cultured in Singapore, Philippines and Indonesia.
Resilience: Medium, minimum population doubling time 1.4–4.4 years (K=0.16-0.20).
Biology and ecology: Occurs in lagoon pinnacles, channels, and outer reef slopes, in coral-rich areas and with
clear waters. Juveniles in seagrass beds. Feeds on fishes, crabs, and cephalopods. May be
ciguatoxic in some areas. Mainly active at dusk.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF J.E. Randall
Epinephelus fuscoguttatus (Forsskål, 1775)
FIGURE 17
Brown-marbled grouper (Epinephelus fuscoguttatus)

Capture-based aquaculture: global overview230
FIGURE 20
Distribution of Epinephelus lanceolatus (FishBase, 2007)
TABLE 10
Characteristics of the giant grouper, Epinephelus lanceolatus
Common names: Giant grouper, Queensland grouper
Size and age: 270 cm TL; max. published weight: 455.0 kg
Environment: Reef-associated; brackish; marine; depth range 1–100 m
Climate: Tropical; 28°N - 39°S, 24°E - 122°W
Importance: Important in subsistence fisheries, commercial aquaculture, recreational gamefish. Cultured
in Taiwan PC. In live reef fish markets. Juveniles sold in ornamental trade as “bumblebee
grouper”.
Resilience: Very low, minimum population doubling time more than 14 years.
Biology and ecology: The largest bony fish found in coral reefs. Common in shallow waters. Found in caves or
wrecks; also in estuaries, from shore and in harbours. Juveniles secretive in reefs and rarely
seen. Feeds on spiny lobsters, fishes, including small sharks and batoids, and juvenile sea
turtles and crustaceans. Nearly wiped out in heavily fished areas. Large individuals may be
ciguatoxic.
Source: Modified from FishBase (Froese and Pauly, 2007).
FAO
Epinephelus lanceolatus (Bloch, 1790)
FIGURE 19
Giant grouper (Epinephelus lanceolatus)

Capture-based aquaculture of groupers 231
FIGURE 22
Distribution of Epinephelus malabaricus (FishBase, 2007)
TABLE 11
Characteristics of the Malabar grouper, Epinephelus malabaricus
Common names: Malabar grouper, estuary grouper, green grouper
Size and age:  234 cm TL; max. published weight: 150.0 kg
Environment: Reef-associated; amphidromous; brackish; marine; depth range 0–150 m
Climate: Tropical; 30°N - 32°S, 29°E - 173°W
Importance: High value commercial and recreational fisheries and aquaculture. Cultured throughout
Asia. Along with E. coioides, the most common species in live reef fish markets.
Resilience: Very low, minimum population doubling time more than 14 years. Listed as Near
Threatened (NT)  by the  IUCN Grouper and Wrasse Specialist Group.
Biology and ecology: A common species found in a variety of habitats: coral and rocky reefs, tide pools,
estuaries, mangrove swamps and sandy or mud bottom from shore to depths of 150 m.
Juveniles found near shore and in estuaries; sex reversal probable. Feeds primarily on fishes
and crustaceans, and occasionally on cephalopods.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF D. First
Epinephelus malabaricus (Bloch and Schneider, 1801)
FIGURE 21
Malabar grouper (Epinephelus malabaricus)

Capture-based aquaculture: global overview232
FIGURE 24
Distribution of Epinephelus quoyanus (FishBase, 2007)
TABLE 12
Characteristics of the longfin grouper, Epinephelus quoyanus
Common names: Longfin grouper
Size and age: 40.0 cm TL
Environment: Reef-associated; marine; depth range 0–50 m
Climate: Tropical; 35°N - 32°S, 110°E - 156°E
Importance: Commercial fisheries and minor aquaculture; in China Hong Kong SAR live fish markets.
Resilience: Medium, minimum population doubling time 1.4–4.4 years.
Biology and ecology: Inhabits inshore silty reefs; there are no records from depths greater than 50 m. Feeds on
crustaceans, fishes, and worms. The enlarged fleshy pectoral fins appear to have resulted
from its habit of sitting on the substrate.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF J.E. Randall
Epinephelus quoyanus (Valenciennes, 1830)
FIGURE 23
Longfin grouper (Epinephelus quoyanus)

Capture-based aquaculture of groupers 233
FIGURE 26
Distribution of Epinephelus sexfasciatus (FishBase, 2007)
TABLE 13
Characteristics of the sixbar grouper, Epinephelus sexfasciatus
Common names: Sixbar grouper, six-banded grouper
Size and age:  40.0 cm TL
Environment: Reef-associated; marine; depth range 10–80 m
Climate: Tropical; 21°N - 21°S, 94°E - 143°E
Importance: Fisheries: commercial.
Resilience: Medium, minimum population doubling time 1.4–4.4 years (K=0.16).
Biology and ecology: Common on silty sand or mud bottoms. Its preference for soft-bottom habitats may
account for its restricted distribution and absence at oceanic islands. Feeds on small fishes
and crustaceans.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF CSIRO
Epinephelus sexfasciatus (Valenciennes, 1828)
FIGURE 25
Sixbar grouper (Epinephelus sexfasciatus)

Capture-based aquaculture: global overview234
FIGURE 28
Distribution of Epinephelus trimaculatus (FishBase, 2007)
TABLE 14
Characteristics of the threespot grouper, Epinephelus trimaculatus
Common names: Threespot grouper
Size and age: 40.0 cm SL
Environment: Reef-associated; marine
Climate: Tropical; 37°N - 20°N, 112°E - 143°E
Importance: Commercial fisheries and minor aquaculture. In China Hong Kong SAR live fish markets.
Resilience: Medium, minimum population doubling time 1.4–4.4 years.
Biology and ecology: Juveniles are common in tide pools and in shallow clear water around rocks and coral
reefs; adults found in deeper water.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF A. Cornish
Epinephelus trimaculatus (Valenciennes, 1828)
FIGURE 27
Threespot grouper (Epinephelus trimaculatus)

Capture-based aquaculture of groupers 235
FIGURE 30
Distribution of Plectropomus leopardus (FishBase, 2007)
TABLE 15
Characteristics of the leopard coralgrouper, Plectropomus leopardus
Common names: Leopard coralgrouper, coral trout
Size and age: 120 cm SL; max. published weight: 23.6 kg; max. reported age: 26 years. On the Great
Barrier Reef, lifespan is 14 years.
Environment: Reef-associated; marine; depth range 3–100 m
Climate: Tropical; 24; 35°N - 30°S, 106°E - 178°W
Importance: Commercial and recreational fisheries and aquaculture, juveniles in ornamental trade.
Resilience: Medium, minimum population doubling time 1.4–4.4 years (tm = 2–4; tmax = 26; Fecundity
= 457  900). Listed as Near Threatened by IUCN Grouper and Wrasse Specialist Group.
Biology and ecology: Inhabit coral-rich areas of lagoon reefs and mid-shelf reefs. Juveniles in shallow water in
reef habitats, especially around coral rubble. Adults piscivorous. Juveniles feed on small fish
and invertebrates such as crustaceans and squid. A protogynous hermaphrodite forming
spawning aggregations on a reef around the new moon.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF J.E. Randall
Plectropomus leopardus (Lacepède, 1802)
FIGURE 29
Leopard coralgrouper (Plectropomus leopardus)

Capture-based aquaculture: global overview236
FIGURE 32
Distribution of Plectropomus maculatus (FishBase, 2007)
TABLE 16
Characteristics of the spotted coralgrouper, Plectropomus maculatus
Common names: Spotted coral grouper, spotted coral trout
Size and age:  100.0 cm SL; max. published weight: 25.0 kg
Environment: Reef-associated; marine; depth range 5–100 m
Climate: Tropical; 21°N - 28°S, 117°E - 159°E
Importance: Commercial and recreational fisheries and aquaculture. In China Hong Kong SAR live fish
markets. Commonly used for food. Its flesh is delicate and well appreciated.
Resilience: Medium, minimum population doubling time 1.4–4.4 years (K=0.21; tm=2–3).
Biology and ecology: Found in protected coastal reef in mixed algae and coral habitat Common on inshore
coastal reefs but absent in clear offshore reefs. Juveniles in shallow water in reef
habitats, especially around coral rubble. Migrate over short distances to spawn, forming
aggregations.
Source: Modified from FishBase (Froese and Pauly, 2007).
COURTESY OF J.E. Randall
Plectropomus maculatus (Bloch, 1790)
FIGURE 31
Spotted coral grouper (Plectropomus maculatus)

Capture-based aquaculture of groupers 237
Trends in production of cultured grouper
Groupers are cultured in many Southeast Asian countries, including Indonesia,
Malaysia, Philippines, Taiwan  Province of China, Thailand, China Hong Kong
SAR, the southeast of China and Viet Nam (Sadovy, 2000). Grouper culture is also
undertaken in India, Sri Lanka, Kingdom of Saudi Arabia, Republic of Korea, Australia,
the Caribbean as well as in the southeastern United States of America.
Despite the huge popularity of live fish in China and Southeast Asia, only
15–20  percent of the amount consumed each year comes from aquaculture, as culture
is principally constrained by limited and unreliable supplies of wild seed and the
difficulties of spawning in captivity. However, hatchery production has increased in
recent years (e.g. Taiwan  Province of China and Kuwait) (Tucker, 1999). It is difficult
to get accurate statistics on farmed grouper production because statistics do not
differentiate between those simply being caught from natural sources and held for a
few weeks in cages before being sold, and those cultured for a longer period of time
(Ottolenghi et al., 2004).
Grouper production through aquaculture is mainly reported by countries in
Asia, where over 9  300 tonnes were produced in 2000. The actual figures of grouper
production in Southeast Asia are reported by Sadovy (2000) to be far higher, at 23  000
tonnes; however, about 20  percent of this production may be based on hatchery
produced fry, while the remainder is from wild seed. Kongkeo and Phillips (2002)
estimated Asian production to be around 15  000 tonnes. In each case, these figures are
significantly higher than the official statistics published by FAO. According to official
statistics, Taiwan Province of China was the leading producer, with nearly 5 100 tonnes
(54 percent of the global total). A total of 7  200 tonnes was produced in brackish water
in Taiwan Province of China, Malaysia and Thailand. The remaining production was
from mariculture a total of 2  100 tonnes, mainly in Indonesia, China Hong Kong SAR
and Taiwan  Province of China.
Grouper culture systems
There are many different systems used for the culture of groupers worldwide,
although there seems to be an agreed set of stages: nursery, transition, and on-growing
(Ottolenghi et al., 2004). Grouper seed must be nursed before being cultured to
marketable size. The nursery stage is reared either in tanks, net cages and hapas
(nylon netting enclosures), or in earthen ponds. Grading is a prerequisite to minimize
cannibalism, especially in the nursery and early grow-out stages. After nurseries, there
are two main systems used for on-growing: pond culture or cage culture. The stocking
density and rearing conditions in both nursery and grow-out phases vary, depending
on the site, the fish sizes, and the grouper species cultured.
Wild fry (2.5–7.2 cm) or fingerlings (7.5–12 cm) may initially be held in tanks or net
cages or earthen ponds for a month or more (nursing period) after harvest (Ottolenghi
et al., 2004). The density may range from 100 to 150 fish/m
2
, e.g. a net of 2  x 2 x 2 m
would hold 400–600 fingerlings. Sorting is undertaken weekly and stock sampling
every 2 weeks. Groupers are normally retained in the nursery until they reach about 16
cm, when they are thinned out and transferred to transition nets (5 x  5 x 5 m) that each
hold 1  100 fish. The fish are finally transferred to production nets after 2–3 months.
Floating cages are often constructed from bamboo poles and polyethylene netting
material (25–50 mm mesh size). Net cages are formed by two types of panels: 4 side
panels forming the walls, and one bottom panel. The net is secured to the raft structure
(bamboo poles) by ropes. Ropes are also used to lash the bamboo poles together.
Buoyancy is provided by empty plastic containers attached to the bamboo frames
(www.seafdeec.org.ph). Net cages come in several sizes (3  x 3 x 2.5 m; 4 x 4 x 2.5 m;
10 x 10 x 3 m); the mesh size ranges from 10 to 35 mm (Agbayani, 2002). The optimum
stocking density averages 120 fish/m
3
. Growth to marketable size (600–800  g) takes

Capture-based aquaculture: global overview238
approximately 8 months, with survival rates of 50  percent or less. Groupers can grow
to 600 g in 12 months, to 1 kg within 18 months, and to 2 kg within 24 months (Tucker,
1999).
Harvesting of groupers is relatively simple (Ottolenghi et al., 2004). Selective
harvesting of groupers weighing 400–600 g is best. A drag net is placed at the farthest
end of the pond or cage, and dragged slowly towards the other end in the early morning.
Fish are then transferred to a holding net where grading is carried out; undersized fish
are returned to the pond or cage.
FISHERIES FOR JUVENILE GROUPER
Collection of grouper seed
Grouper seed is collected using several different methods, depending on location
(Table 17). Capture methods are generally artisanal and the fishermen employ a variety
of artificial habitats. Moreover, different fishing gears are used at different times of the
year: the gear change follows the growth of the seed and their movement to deeper
waters as the season progresses. Gears used to take grouper seed can be divided into 8
different categories: large fixed nets (e.g. fyke nets), traps and shelters, hook and line,
scoop and push nets, artificial reefs, fish attractors, tidal pools and chemicals. The sizes
of grouper seed caught and traded vary between 1 and 25 cm, i.e. from the moment of
settlement to fish that are over one year old. However, most of the catch focuses on
fish up to about 15 cm (Sadovy, 2000).
Some grouper seed collection methods are more damaging than others. Clearly
destructive methods include those that result in high mortality, involve high levels of
TABLE 17
Overview of seed collection methods for capture-based aquaculture of groupers in Southeast Asia
Gear type Description Location Fish size (cm)
Gango (fish
nests)
Conical pile of waterlogged, criss-crossed wood or of
rocks, sometimes in combination, together with old car
tires, PVC pipe cuttings, bamboo sections, or other shelter
materials. Covers 5–10  m², with a 2–3  m diameter or
2.5–3x2–3  m base and 0.5–1.5  m height. The largest may
be 5 m diameter at the base.
Philippines 2–15 cm
Fish shelters Formed by hanging brushes, nets or clusters of grasses,
leaves or other materials. Used with or without lights.
Philippines
China
Thailand
1–3 cm
Fish traps Vary in shape and size, and in mesh size. The trap frame is
made of metal, wood or bamboo.
Indonesia
Malaysia
Philippines
China
Taiwan PC
Viet Nam
2–25 cm
Fyke netBig collectors, stationary nets installed in river mouths
during high tides. Three mesh sizes are used: larger at the
aperture, followed by medium and finer net at the end.
Philippines
Thailand
Viet Nam
1–15 cm
Hook and line  Indonesia
Malaysia
Philippines
China
Taiwan PC
Thailand
Viet Nam
>7.5 cm
Scissor net A triangular bamboo frame of various dimensions, which
may or may not have “shoes” to assist it in moving over
the substrate. Fine meshed netting is attached to the
frame and the bamboo poles are crossed over each other.
Philippines
Thailand
2.5–15 cm
Miracle hole Shallow holes are excavated on tidal flats. Sometimes the
wall of the hole is built up with rocks.
Philippines 5–10 cm
Temarang Artificial aggregating device (fish shelter), which consists
of a bunch of twigs from wild shrubs; about 20–30
bunches of 50 cm length are tied to a 5 m rope and hung
over sandy sea bottom between two poles.
Malaysia 2–2.5 cm
Source: Modified from Ottolenghi et al., 2004.

Capture-based aquaculture of groupers 239
bycatch, cause damage to the fish habitat and/or result in monopolization of the local
fishery by a few individuals. Destructive methods include scissor nets and fyke nets,
which are already banned in some areas. Lift nets are also destructive, particularly
in terms of bycatch. Gangos, miracle holes and other types of artificial shelters and
seed aggregation devices do not possess the above drawbacks. Methods that target
postlarvae seem less likely to deplete wild stocks because of the high natural mortality
that probably characterizes this stage in the wild (Johannes and Ogburn, 1999; Sadovy,
2000).
Mortality rates from catching to stocking
Seed quality depends on the type of fishing gears used, and there are significant
differences in seed mortality rates. Mortality rates associated with fish traps are usually
low. For example, the use of “Bubu” (fish traps used in Malaysia) cause a 5  percent
mortality rate, while artificial aggregators such as Temarang (also used in Malaysia)
cause 3  percent mortality. Other catching methods, like scissor nets and fyke nets,
can generate a high mortality. “Pompang” (fyke net) and “Wunron” (push/scissor
net), which are used in Thailand, are reported to cause 20–30  percent and 80  percent
mortality rates, respectively (Sadovy, 2000). It is likely that subsequent mortalities
during transport and stocking will also be high, as many of the seed fish will also have
been damaged, and are therefore susceptible to stress and disease.
The problems that arise during seed transport to the net cages or to the middleman/
farmer/exporter, depend on seed size, quality, fitness and the locality. For transport
over short distances, in Thailand, for example, “seeds” are placed in styrofoam boxes
or buckets, with or without aeration (often provided by middlemen), or with holes in
the bottom for water exchange.
Transport time is typically from about 10 minutes up to two hours. Post-harvest
mortality is low. For longer transit periods, fish are packed in 23–25°C seawater with
aeration. Transport densities are about fifty 7.5 cm fish per bag, or one hundred 1 cm
fish/l, or two to three hundred 3–7.5 cm fish per bucket. For a 7-hour journey, ice can
be used to keep the water cool. Some exporters use an anaesthetic, either quinaldine
or MS222, but consider the latter to be rather expensive. The use of anaesthetic is
considered important to reduce the likelihood of spines piercing the plastic transport
bag. For export, fish are packed into styrofoam boxes of various sizes; each shipment
has about 20 000 fish in 30 boxes (Sadovy, 2000).
In the Philippines, approximately 10  percent of the seed caught is used domestically,
while the remainder is exported. There can be significant mortalities during local
transportation. The movement of seed from the catchers to the middlemen or the
farmers is carried out by keeping fish in plastic containers or basins with holes for
water circulation. Mortality rates are quite low under such circumstances. If destined
for trade, the fish may be maintained for short periods by the middleman, prior to
packing and shipping, either domestically or internationally. In some cases, they may
be transferred temporarily (for a few days) to an “aquarium box” to await buyers
who come to collect fish and who are responsible for the export of the fry. Mortality
rates can reach 10–20  percent at this stage, i.e. prior to selling to buyers for export or
domestic trade (Sadovy, 2000). Mortality rates are low if the transit time is less than an
hour. However, for longer periods, if there is no aeration or frequent water changes,
mortality increases and oxygen may have to be added. Buyers pack fry in double plastic
bags with pre-cooled water using ice (18–22°C) and a salinity of 15–18 ppt. 2.5 cm
“seeds” are packed 400–500 per box and 7.5–10 cm “seeds” are packed 20–40 per box
(Sadovy, 2000).
The mortality rates that follow capture and transport are not exactly known;
estimates for over the first 2 months after catching are quite variable (30–70  percent),
depending on the quality of fry, the level of transport stress, and the presence

Capture-based aquaculture: global overview240
of disease and cannibalism (Pudadera, Hamid and Yusof, 2002). According to a
report from the Secretariat of the Pacific Community (www.spc.org.nc/coastfish/
News/LRF/5/15GrouperHK.htm), the survival rate for imported fry is low, at
10–20  percent.
AQUACULTURE DEPENDENCY ON WILD SEED
Generally, groupers spawn on offshore reefs where they form aggregations of hundreds
to tens of thousands of individuals, in a few specific locations (Johannes et al., 1999;
Rhodes and Sadovy, 2002). They produce pelagic larvae that may disperse over hundreds
of kilometres in the course of 30–45 days and experience high density-independent
mortality. However, recent research suggests that groupers and other reef fishes may
have greater control over their distribution than previously thought, and that at least
some proportion of the gametes spawned may be retained near their natal reef (Jones
et al., 1999; Jones, Planes and Thorrold, 2005; Swearer et al., 1999). Larvae, transported
to near-shore nursery habitats settle as juveniles in sea-grass beds, mangroves, algal
beds, coral rubble, oyster reefs and marshes (Coleman et al., 1999; Tupper, 2007). For
this reason grouper seed is mainly caught in coastal areas, particularly around sea-grass,
mangrove and shallow brackish water areas near river mouths and estuaries, as well as
in tidal pools, tidal channels and around reefs.
The peak grouper seed season is often associated with the relatively wet months in
the year (e.g. monsoon seasons); in several areas, grouper seed collectors have claimed
that their best catches were associated with strong onshore winds (Johannes and
Ogburn, 1999). This is consistent with a number of recent studies into recruitment
pulses of settlement-stage reef fish – including groupers – that accompanied cyclonic
storms, which apparently caused the fish to be transported shoreward (Shenker et al.,
1993; Dixon, Millich and Sugihara, 1999).
Because grouper are particularly difficult to culture in closed systems, full-cycle
culture of most grouper species is not yet possible. For this reason, approximately
66–80  percent of all grouper culture involves the capture and grow-out of wild seed
(Sadovy, 2000). The volume of seed caught each year exceeds hundreds of millions of
individuals (Sadovy, 2000). The greatest catches tend to be of the smallest size classes
(1–3 cm); during peak seasons a catch can be of tens of thousands by a single unit of
gear, in a single night, by one fisherman (e.g. using a fyke net). Even larger sizes of fish
are being captured in massive numbers region-wide each year. It is important to realize
that the equivalent of the typical annual amount of seed produced in the hatcheries in
the whole of Southeast Asia (excluding Taiwan  Province of China), i.e. 20  000 to 80  000
fry, can be caught by one fisherman in one night (Sadovy, 2000).
When seed catches are compared to the numbers of marketable fish produced, the
results strongly suggest crude and wasteful culture practices. Sadovy (2000) estimated
that about 60 million seed fish are needed produce the regional total of 23  000 metric
tonnes of table-size live fish from culture annually.
FISH FEED
As with all culture systems, there are many local variations in the feeds and feeding
regimes utilized. There appears to be no universal system, and local availability seems
to be the key criteria in developing a feeding schedule (Ottolenghi et al., 2004). Fry and
fingerlings are fed with mysids and small shrimp for a couple of days post-capture in
tanks, to acclimatize them and check that all individuals are eating. Trash fish forms the
main feed in nursery and production cages, which is minced or chopped to suit each
size group; trash fish may be supplemented with vitamins and minerals. This kind of
feed is gradually being replaced by moist pelleted feed.
Trash fish is commonly used for feeding in grouper cage culture, but its increasing
cost, shortage of supply, variable quality and poor feed conversion ratios indicate that

Capture-based aquaculture of groupers 241
this form of feed may not be the best from either a nutritional or an economic point of
view. However, groupers fed with bycatch (trash fish) in a study by Bombeo-Tuburan,
Kanchanakhan and China (2001) fared significantly better in terms of final length and
total production than when fed other diets (live tilapia, formulated diet).
A major problem is the limited supply of trash fish, so there is a need to develop a
suitable diet for grow-out grouper production (Millamena, 2002). Fishery products, both
in the form of low value trash fish or fishmeal, are presently the major sources of protein
in the grow-out culture of most fish species and constitute up to 70  percent of their
dietary composition. As the demand for fishmeal and fish oil for aquaculture increases,
costs are expected to rise unless new sources (e.g. fish discards, krill, mesopelagics) can
be economically exploited or substitutes for these marine products for inclusion in
aquafeeds prove commercially applicable (New and Wijkstrom, 2002).
A dependable supply of cost-effective, non-marine, sources of alternative protein
must be provided if fish farming is to remain profitable. Millamena (2002) conducted
a feeding trial to evaluate the potential of replacing fishmeal with processed animal
by-product meals, meat meal and blood meal, in practical diets for juvenile groupers
(Epinephelus coioides). The study demonstrated that up to 80  percent of fishmeal protein
can be replaced by processed meat and blood meal derived from terrestrial animals with
no adverse effects on growth, survival, and food conversion ratio (FCR). From an
economic standpoint, replacement of fishmeal with cheaper animal by-product meals in
practical diets can alleviate the problem of low fishmeal availability and high costs. These
processed by-products can be delivered in the Philippines, for example, at US$0.40/kg,
less than half the price of most commercial fishmeals (US$1/kg). The effective use of
meat meal-based diets for grouper grow-out also reduces the requirements for trash
fish, another fishery resource that is extensively used (Millamena, 2002). Economic
sensitivity analysis showed that a combination of improvements resulted in higher
return-on-investment (ROI). However, these apparently favourable results must be
balanced with the fact that some countries (e.g. in the EU) have banned the inclusion of
all terrestrial meat-meal based products in fish feeds, due to fears concerning mad-cow
disease (Ottolenghi et al., 2004).
ENVIRONMENTAL IMPACTS OF THE JUVENILE GROUPER FISHERY
Mous et al. (2006) conducted a pilot study in Indonesia of artificial shelters (gangos), to
determine the sizes and capture rates of species of interest to the live fish trade, and to
determine the likely environmental footprint of a gango type of capture method. From
the results of the 15-month study, they drew inferences regarding the sustainability of
this fishing method and requirements of space, fish and materials for a viable grow-out
operation. The results showed that gangos were unselective for either species or size.
Only 1.4  percent of the total fish catch (by number) were target species, mainly the
grouper Epinephelus coioides, and most were large enough (mean total length was 13.6
cm) to have bypassed the early high mortality phase. Moreover, there were large non-
target catches that included many food fish species too small to be useful in catches.
Assuming that a soak-time of 3 months results in an average catch per gango of 6.6
E. coioides (as was observed for this species in Terang Bay, the most productive of
the four sites), yearly production per gango would amount to 26.4 fish. Even a small
local grow-out industry with a capacity of 25 tonnes would require an annual supply
of 80  000 fish, assuming a grow-out weight of 0.6 kilogram and 50  percent mortality
from fingerling stage to market-ready product. This would require deployment of an
estimated minimum of 3  000 gangos.
With such figures in mind, Mous et al. (2006) estimated of the space needed to
accommodate sufficient gango deployment and suggested that a sizeable fish culture
industry based on capture of fry, fingerlings and juveniles from the wild would have
a large ecological footprint. For example, the 3  000 gangos estimated to support a

Capture-based aquaculture: global overview242
25 tonnes grow-out operation, would require approximately 300  000 m
2
(assuming that
each gango requires a plot of 10  x 10 m) or 30 kilometres of coastline (assuming that
gangos are deployed in a single line following the optimum depth contour). In other
words, juvenile supply would require 1.2  hectares of shallow coastal waters for each
tonne produced.
Given the large number and area of gangos needed for a viable operation, and that many
groupers captured could probably have survived to reproduce, the ecological footprint
of this approach could be substantial (Mous et al., 2006). These results, and literature on
other juvenile fisheries, suggest that CBA sources of seed such as gangos may often need
management, have important links to other capture fishery sectors, and require careful
evaluation of potential costs and benefits before introduction or development.
Support for grouper CBA is often based on the assumption that the natural morality
of early juvenile grouper is very high, so that the fishery is not adding substantially to
this natural mortality and therefore not affecting adult population size to any great
extent. This assumption remains untested for grouper species. As Sadovy (2001b)
points out, the critical question is how early do juvenile mortality rates decline to
adult levels? If early mortality is high, then removal of some post-settlement fish for
culture may have little impact on adult numbers, since the probability of survival of
any individual fish is low. However, recent research suggests that the period of very
high mortality occurs during and immediately after settlement, and that fish surviving
more than a few days have a much higher chance of survival. Tupper (2007) estimated
the cumulative mortality of early juvenile (2.5–5.0  cm TL) Plectropomus areolatus
and Epinephelus polyphekadion in their preferred nursery habitats to be around
50–75  percent over the first 3 months post-settlement. Assuming an exponential rate
of decline in mortality, the instantaneous mortality at 3 months post-settlement would
be much lower than 50  percent. Indeed, mortality rates of post-settlement juveniles
may not be substantially greater than adult mortality (estimated at 20–30  percent for
most groupers, e.g. Posada and Appledoorn, 1996) and are likely much lower than the
estimates of >90  percent mortality often suggested for newly settled reef fishes. If each
individual has a 50  percent chance of surviving the first 3 months after settlement, then
removal of large numbers of juveniles will almost certainly have an impact on adult
population size. This could result in direct conflicts with the adult capture fishery and
could accelerate overfishing of groupers.
In addition to problems of bycatch, wasteful mortality, and overfishing, cage and
net culture can create other environmental problems, most notably point-source
pollution which can have adverse effects on coastal waters, and particularly on coral
reefs. For example, in 1994, researchers in Barbados noted complete bleaching and
eventual death of coral patch reefs in the vicinity of a cage culture operation for dolphin
fish (Coryphaena hippurus). Disease transfer is another problem exacerbated by the
complex and extensive trade in live fish between Asian countries.
SOCIAL AND ECONOMIC IMPACTS OF GROUPER FARMING
Despite the growing importance of grouper aquaculture as both an alternative to wild
caught grouper for the LRFT, and as an alternative livelihood for fishers engaged in
destructive fishing practices, relatively little is known about the social and economic
impacts of grouper farming, and the broader socio-economic context in which it takes
place. Studies have focused on the trade of live reef fish which fuels the fishery for
grouper and provides an incentive for grouper aquaculture.
The trade in live reef fish
The trade in live reef fish, whereby fish are transported live from the capture location to
restaurants and supermarkets, began in China in the 1960s when a few marine species
were to be found in the live fish markets of China Hong Kong SAR, and has expanded

Capture-based aquaculture of groupers 243
rapidly since the early 1990s. The preference for keeping fish alive until minutes before
cooking and consumption has been popular for centuries in Chinese culture, and until
recently this demand for live fish was supplied by locally caught species. A preferred
species for consumption was the red grouper, Epinephalus akaara, until overfishing
of both adults and later fingerlings for culture in China Hong Kong SAR waters led
to severe depletion of local stocks, forcing fishermen and the LRFT industry further
afield to seek out supplies to meet local demand for market size fish. In the mid-1970s
fishing boats began to exploit Philippine waters, and later the islands of Indonesia,
before moving on to the Pacific Islands (e.g. Papua New Guinea, the Solomon Islands),
Australia’s Great Barrier Reef, and the Maldives (Johannes and Riepen, 1995). Thailand
is now also an important contributor to the LRFT. The trade supplies a luxury, niche
market. Live reef fish are described as being “high-value-to-volume” and can fetch
US$5 to US$180 per kilogram, considerably more than dead reef fish (Sadovy et al.,
2003). Highly valued species such as Cheilinus undulatus, or humphead wrasse, can
fetch a price of up to US$200 per kilogram (Lau and Parry-Jones, 1999).
China Hong Kong SAR is the hub of the live reef fish trade, and the destination for
much of the wild-caught and cultured grouper in the region. Approximately 60  percent
of internationally traded live reef fish are exported to China Hong Kong SAR (Sadovy
and Vincent, 2002), representing approximately 15  000 to 20  000 tonnes per year at a
value of US$350 million (Muldoon and MacGilvray, 2004). Accurate volumes of trade
for individual species are difficult to estimate, as exports are not disaggregated at the
species level and much of the trade goes unreported (Sadovy et al., 2003).
The market network linking farmers to consumers is relatively long and complex,
frequently crossing international boundaries, with ownership changing repeatedly.
Grouper farmers obtain fry fish from their own fish catch, purchase from local fry
fishers, private or government hatcheries. It is common for fry fishers who do culture
grouper to sell their catch to a middleman, who may support a group of ten to thirty
fishers. The fry are then either sold locally to farmers for on-growing or transported
directly to export centres for shipping to other countries in the region. Grouper from
grow-out operations are also predominantly destined for the export market, although
there is also a growing domestic market in many countries where grouper are becoming
increasingly popular on the menus of local seafood restaurants throughout Southeast
Asia.
Social impact of grouper fry fishing
The number of fishers exploiting the grouper fry resource is unknown, but estimates
suggest that fry fishers in the Philippines may number in the tens of thousands (Sadovy,
2000). For these fishers, fry fishing represents one activity in a broader portfolio of
activities on which they depend. Fry fishing is seasonal in nature and both fishers and
non-fishers alike enter the fry fishery if market signals indicate a lucrative opportunity.
Fishers may be engaged in the fishery on a full- or part-time basis, whilst also engaging
in other fishing activities for the capture of food fish or fish for the aquarium trade
(Sadovy, 2000). The capture of wild grouper fry is reported to make a significant
economic contribution to the lives of coastal fishers (Sadovy, 2000). However, despite
this apparent significance, few studies have attempted to assess the role of these wild
fry fisheries in the livelihoods of coastal fishers. There is, therefore, a critical gap in our
understanding of the precise nature of the contribution made by wild fry fisheries to
coastal households, the economic and gender profile of fishers, and the way in which
coastal fishers may be affected by developments in the grouper industry.
Some studies in the region do indicate that the capture of grouper fry may contribute
substantially to household incomes. In Sulawesi, Indonesia, for example, fishers may
catch in the region of 1  000–2  000 2.5 cm fry per fisher on daily basis during the peak
season using scoop nets, with a value of US$300–600 (Haylor et al., 2003). In Viet

Capture-based aquaculture: global overview244
Nam, income from grouper fry/fingerling harvest was reported to earn fishers as much
as US$3  080 per year (Sadovy, 2000).
Grouper fry fishers do not represent a homogenous group in terms of social status.
Fishing households, like most rural households, engage in a diverse range of activities of
which fishing may be only one component. Similarly, fishing activities are also diverse
with fishers using a variety of gears to target different species according to seasonality
and the tides. Dependence upon fry fishing is therefore rare, if it exists at all, although
the extent to which the income from fry capture contributes to the total household
income will vary from household to household. The relatively high value of grouper
fry compared to the rest of the fish catch may, therefore, represent an important income
source. As one fisher in Viet Nam indicated, catching 5–10 grouper fry per day can equal
the income from all the other fish harvested (Sadovy, 2000). Findings from a survey in
Thailand suggest that, for the majority of households, fry fishing is a supplementary
activity, often opportunistic, with fishers entering and leaving the fishery according to
fry abundance and market signals. Fry fishing in southern Thailand complements the
regular fishing activities of coastal fishers, whose principal target species, including
shrimp, small pelagic species, are caught at different times of the lunar calendar. Fry
fishing therefore allows fishers to supplement their fishing activities at a time when
fishing would otherwise not be possible (Sheriff, 2004).
Social impacts of grouper aquaculture
Important synergies exist between grouper aquaculture and fry fishing, which blur the
distinction between fry fishers and grouper farmers, and give added significance to the
role of grouper fry in coastal livelihoods. In the absence of a reliable source of hatchery
fry, and the preference of many farmers for wild caught fry even where hatchery fry
is available, most grouper farmers rely upon wild-caught fry to stock their culture
systems. Where adequate supplies of grouper fry are still available in the wild, many
farmers fish for their own seed inputs which, as they are not purchased and require no
cash outlay, are considered a “free” resource. This has important implications for the
ability of resource poor households, with little access to financial capital, to take up
grouper aquaculture.
Grouper aquaculture has been identified as an activity which can generate a relatively
high return in comparison to many of activities available to coastal households
(Hambrey, Tuan and Thuong, 2001). Many activities which generate a comparable
return, including trading, ownership of plantations and shrimp culture are inaccessible
to the majority of households due to the high levels of investment required to take
up these activities. Grouper aquaculture can therefore be an important addition to
household livelihoods, providing a means of savings to supplement the daily income
generated by regular fishing activities (Sheriff, 2004).
As a solution to the problem of destructive fishing, aquaculture may not present
the ideal alternative to fishing, as is frequently suggested. There is an assumption that
aquaculture is an activity that is easily interchangeable with fishing as a livelihood
activity, and that fishers are willing and able to give up fishing to take up a new and
markedly different occupation. Studies suggest that fishing is deeply rooted in the lives
and traditions of “fishing” communities and the identity of fishers. McGoodwin (2001)
reports that fishing is regarded “not merely as a means of ensuring their livelihoods, but
as an intrinsically rewarding activity in its own right – as a desirable and meaningful way
of spending one’s life…prompting many fishers to tenaciously adhere to the occupation
and to continue fishing even after it has become economically unrewarding.” In a study
conducted by Pollnac, Pomeroy and Harkes (2001) in the Philippines, Indonesia and
Malaysia, it was found that, in all three countries, fishers like their occupation and
only a minority would change to another occupation, with a similar income, if it were
available. In the Philippines, 95  percent of fishers surveyed reported that they would

Capture-based aquaculture of groupers 245
choose to become fishers again if they had to live their life over again. They also cited
pleasurable aspects of the job as reasons for staying in the fishery, including the beauty
of the sea and not having to work for a boss. Fishers in the three countries who would
choose to leave the fishery were characterized by a higher level of education and a lower
income from fishing. The results do not support the view that fishers are the poorest of
the poor, as fishers cite income as one of the reasons for choosing not to change their
occupation. The level of satisfaction with fishing as an occupation suggests that fishers
will not necessarily change to an alternative occupation and leave the fishery (Pollnac,
Pomeroy and Harkes, 2001). Furthermore, the role of fishing in households livelihoods
differs markedly from the contribution made by aquaculture. Fishing provides a source
of daily income which pays for the daily needs of the household. In contrast, fish culture
has been identified as being of importance to the households ability to save money
and thus to accumulate assets. Proposals to encourage fishers to leave the fishery by
offering fish culture as an alternative, may therefore fail, as fish culture cannot meet
the daily needs of the household. Aquaculture can, however, provide an important
supplementary activity to support livelihood diversification in coastal communities,
where few alternatives may exist (Sheriff, 2004).
As a contributor to rural livelihoods, particularly those of coastal fishers, grouper
aquaculture can generate potentially large financial benefits. The high-value of grouper
on the export market ensures that farmers are able to generate a profit even when stocks
suffer heavy mortalities. High initial investment cost is frequently cited as the principal
constraint to the uptake of grouper aquaculture. Approximate investment costs for
a small-scale farm are in the region of US$1  470 in the Philippines and US$1  010 in
Indonesia (Pomeroy, Parks and Balboa, 2006), US$516 in Viet Nam (Hambrey, Tuan
and Thuong, 2001) and US$237 in Thailand (Sheriff, 2004). Financial analyses of
grouper aquaculture have indicated that grouper rearing is financially feasible, although
Pomeroy, Parks and Balboa (2006) found that the capital requirements of some
aquaculture systems in the Philippines and Indonesia may be beyond the financial
means of many small producers, specifically broodstock and nursery/hatchery
systems (Table 18). However, capital costs for grow-out are substantially lower than
the broodstock or nursery systems, and are within the financial means of many small
producers (Pomeroy, Parks and Balboa, 2006) (Table 19). This figure excludes holding
tanks, and therefore more accurately reflects reality. Fish are most frequently kept in
holding tanks of a local fish trader within the community, and therefore represent a
cost which will not often be incurred at the farm level. The total production costs per
market size fish from grow-out, US$3.01 in the Philippines and US$3.18 in Indonesia
for a 600  g fish, were found to be well below the average selling price at the time of
the Pomeroy, Parks and Balboa (2006) study, which was US$6 in 2002. With the sale
of market size fish able to generate this level of profit, it is not surprising to find that
the annual enterprise budget show in Table 20 suggests that the cost of investment can
be recouped relatively quickly. Pomeroy, Parks and Balboa (2006) conclude from their
analysis that loans or other incentives to cover start-up costs could be repaid within the
first or second year of production.
Despite these apparently high costs studies have shown that, with appropriate
support, even the poorest can benefit from grouper culture, with implications for
both household well-being and community development. For example, one study
in Thailand found that grouper culture was taken up by households from all wealth
groups within a community in Satun province (Sheriff et al., in press). Support from
the Thai Department of Fisheries in the form of materials for cage construction and
seabass seed allowed households to establish a small farm of two cages with which to
initiate grouper culture. Grouper fry were supplied in small quantities from the farmers
fish catch, and a share of the profits returned to a centralized revolving fund for the
benefit of all households in the community.

Capture-based aquaculture: global overview246
The absence of large-scale production of grouper fry has ensured that production is
kept primarily in the hands of small-scale, individual family owned operations (Hambrey
et al., 2001; Sadovy, 2000; Sheriff, 2004), however some systems involve a large number
TABLE 18
Total projected capital investment costs for grouper hatchery/nursery and broodstock operations, modelled
for Indonesia and Philippines (in US$)
Nursery/hatchery operations Broodstock system
Philippines Indonesia Indonesia Philippines Indonesia
(small size
operation)
(medium size
operation)
Broodstock costs
Male specimens 400 660
Female specimens 600 1 320
Sub-total 1 000 1 980
Land operations
Land 6 500 728 4 850 6 500 4 850
Perimeter fencing 50 20 50 50 50
Tanks and reservoirs 11 900 1 110 8 760 6 100 3 080
Roofing, framing and siding 3  800 230 1 835 2 400 1 000
Building/structures 5 800 250 1 850 0 650
Plumbing 4 000 210 2 000 2 000 1 000
Electrical 3 500 135 1 700 2 000 1 000
Air blower 1 980 100 600 1 320 250
Sea- and freshwater pumps 2 600 315 1 800 1 700 900
Generator 2 000 0 450 2 000 450
Truck 0 0 14 000 0 0
Miscellaneous 1 000 160 900 200 150
Sub-total 24 270 13 380
Total capital investment 43 130 3 258 38 795 25 270 15 360
Source: Adapted from Pomeroy, Parks and Balboa, 2006.
TABLE 19
Total projected capital investment costs for grow-out of grouper modelled for Indonesia and Philippines
(in US$)
Indonesia
(Cromileptes altivelis)
Philippines
(Epinephalus coioides and E. malabaricus)
Marine Operations
Floating net cages 1 280 825
Boat 80 60
Water quality test equipment 70 80
Harvest equipment 40 45
Sub-total 1 470 1 010
Source: Adapted from Pomeroy, Parks and Balboa, 2006.
TABLE 20
Summary of annual enterprise budgets over a single production cycle across grouper scenarios modelled for
the Philippines (Epinephalus spp.; 12 month grow-out period) and Indonesia (Cromileptes altivelis; 18 month
grow-out period)
Philippines Indonesia
Broodstock Hatchery /
nursery
Grow-out Broodstock Hatchery /
nursery
(medium)
Hatchery /
nursery
(small)
Grow-out
Variable costs
1
13 128 15  895 4  786 8  160 24  325 3  435 8  363
Fixed costs
2
4 053 7  713 2  299 2  662 8  235 668 2  108
Total expenses 17  181 23  608 7  085 10  882 32  710 4  103 10  471
Total income 23  503 45  045 14  400 83  784 106  270 12  505 20  250
Balance 6 322 21  347 7  315 72  902 73  560 8 402 9 779
Source: Adapted from Pomeroy, Parks and Balboa, 2006.
1
Eggs, fingerlings, feed, vitamins/medication, chemicals, electricity, labour and consultants, fuel and oil, marketing/packing/
harvesting, supplies, repairs.
2
Depreciation of fixed assets, interest payments.

Capture-based aquaculture of groupers 247
of cages and off-shore systems are being tested in countries including Malaysia and
Viet Nam (Kongkeo and Philipps, 2001). On-going research efforts are focusing on
the hatchery production of the most vulnerable and commercially important grouper
species in an attempt to reduce pressure on wild stocks. Yet there may be significant
socio-economic impacts if hatchery production becomes commercially viable on a
large-scale, and may threaten the livelihoods of both fry fishers and small scale grouper
farmers. Taiwan Province of China is one of few countries to have a successful hatchery
industry and may provide some insight into the potential impacts of hatchery produced
grouper, where production has led to a marked effect on demand for grouper fry and a
subsequent decline in seed prices. A reduction in the value of grouper is anticipated by
exporters and importers as a result of increased production (Sadovy, 2001a). However,
small-scale hatchery production of grouper has been found to be a viable livelihood
option providing employment opportunities and rural livelihood diversification (Siar,
Johnston and Sim, 2002). In Bali, where many such hatcheries have been established,
milkfish fry production has provided the basis for diversification into grouper fry,
and therefore provides a particularly relevant model for transfer to countries like the
Philippines. However, uncertainties remain as to the acceptability of hatchery produced
grouper fry to grouper farmers and the likely livelihood impact of hatchery production
on fry fishers and the value of cultured grouper.
Gender roles in the grouper fry fishery and aquaculture
The specific role of women within the grouper fry capture fishery and trade network
is little understood. Within the fisheries sector, women often play an important role
in post-harvest activities, which are absent from the live fish trade. However, women
frequently take responsibility for trade and financial matters, and in countries such as
Thailand, it is not unusual to find that the main fish and fry trader within the community
is female, although fish trading beyond the community is more frequently the domain
of men (Sheriff, personal communication). Grouper culture can provide perhaps the
most significant opportunities for women, who are often responsible for maintaining
aquaculture operations on a daily basis (Haylor et al., 2003). The requirement for
trash fish is high in grouper culture, and the preparation of trash fish for feeding is
frequently done by women (Sheriff, 2004). Experience in Indonesia has shown that
women may also find employment in the small-scale hatchery industry, providing
labour as temporary workers for the counting and packaging of milkfish fry (earning in
the region of US$0.33 per 5  000 fry counted), or as brokers in the fry marketing chain
(Siar, Johnston and Sim, 2002). Similar work in a grouper hatchery grading grouper fry
may earn women US$6.66 per day. The work is however, extremely hard, according to
one hatchery owner (Siar, Johnston and Sim, 2002).
MANAGING CAPTURE-BASED AQUACULTURE OF GROUPER
The management of capture-based farmed groupers is complicated by several problems,
including shortage of capture-based seed, disease transfer resulting from international
trade in seed, high mortality rates in capture and culture, overfishing of grouper adults,
etc. (Ottolenghi  et al., 2004). Groupers are top predators, sedentary in character and
strongly territorial, typically long-lived and slow growing and many assemble in
large numbers to spawn. These characteristics contribute to the ease with which over-
exploitation may occur, and is engendered by the Live Reef Fish Trade (LRFT). This
has already led to calls to include many of the target species in Appendix II or III of the
Convention on International Trade in Endangered Species (CITES) (Lau and Parry-
Jones, 1999). The Nature Conservancy (TNC) has developed a regional strategy in the
Asia-Pacific that focuses on developing and applying regional models to sustainable
fisheries. Many different approaches have been taken to reduce exploitation, e.g. the
Bahamian government has recently approved the establishment of five no-take marine

Capture-based aquaculture: global overview248
reserves. All of these sites contain known Nassau grouper (Epinephelus striatus)
spawning aggregations. Although stocks of Nassau grouper in the Bahamas appear to
be healthy, these closures (coupled with other research activities) are being implemented
to ensure that conservative management measures are taken, as a precaution against
stock collapses such as those that have occurred in other locations that once held stocks
of this species (Johannes, 2000). In Micronesia, Palau was among the first nations in the
world to protect their grouper spawning aggregations, enforcing a seasonal closure on
the Ngerumekaol (Ulong Channel) aggregation site in 1976, then creating permanent
no-take marine reserves at Ngerumekaol and Ebiil (another aggregation site) in 1999.
Pohnpei State in the Federated States of Micronesia has also declared permanent
no-take zones around its grouper spawning aggregations (Rhodes and Tupper, 2007).
Both Palau and Pohnpei have closed their grouper fisheries during the reproductive
season and have limited or banned export of groupers and other species involved in
the live reef fish trade.
Other regulations should be developed to control the harvest of grouper seed. The
availability of capture-based grouper seed is often insufficient and unreliable (both in
quality and quantity) to meet demand; low production in farming is mainly attributed
to lack of seed supply (Chao and Chou, 1999; Yashiro, Vatanakul and Panichsuke,
2002; Agbayani, 2002). Disease problems due to the high transfer stress can cause high
mortality rates in capture and culture. Sadovy (2000) has compiled information on the
status of regulations on grouper seed capture and exports that concern capture-based
aquaculture (Table 21).
A survey of CBA in Southeast Asia found that while the quantity of seed caught
was significant, the production level was very low (Sadovy, 2000). The major causes
contributing to this massive mortality are destructive fishing practices and gears, poor
post-harvest handling, poor farming practices and conditions, and a generalized lack
of experience or knowledge. This review indicated that there is a substantial fishery,
and demand, for fish in the 5–10 cm range, but that the removal of this seed could
have serious consequences for the future of both adult stocks and the contribution
of these adults to the future of the seed fishery itself. Given the likelihood that there
will be a significant increase in natural mortality for the smallest settling fish, several
researchers have already proposed that fisheries for very early post-settlement (or even
pre-settlement) seed is a way of gaining benefit from a resource that does not affect its
long-term sustainability.
It is necessary to consider further initiatives to attain a more sustainable use of grouper
stocks and greater socio-economic benefits from grouper capture-based aquaculture.
One possible approach for grouper management is, as Sadovy (2000) suggests, the
establishment of nursery areas where the capture fishery and culture operations occur.
Another possibility is to protect key seed settlement areas and nursery habitats, such as
mangrove areas, coral rubble and sea-grass environments in river mouths and estuaries,
and to ensure seed production by safeguarding spawning adults. Marine protected areas
(MPAs) should incorporate key settlement and nursery areas, but to date, there are few
(if any) MPAs protecting grouper nursery habitat (Tupper, 2007).
Positive steps to address many of these issues are being taken by the Network of
Aquaculture Centres in Asia and the Pacific (NACA) and its partners, the Asia-Pacific
Economic Cooperation (APEC), the South-East Asian Fisheries Development Center
(SEAFDEC), the Australian Centre for International Agricultural Research (ACIAR),
and the WorldFish Center (formerly known as ICLARM), etc. In 1998 the Asia-Pacific
Grouper Network (APGN) was established; this organization addresses aquaculture
development, in order to:
• reduce the current reliance on capture-based “seed” for aquaculture, as the capture
of wild juveniles is sometimes carried out using destructive fishing techniques that
can have significant impact on the long-term status of the stock;

Capture-based aquaculture of groupers 249
• provide an alternative source of income/employment for coastal populations
currently engaging in destructive fishing practices;
• protect endangered reef fish from the pressures of illegal fishing practices, through
the development of sustainable aquaculture; and
• develop new aquaculture livelihood options and investments that will generate
economic benefits for a diversity of stakeholders and employees.
Since 1996, all the above mentioned organizations have conducted workshops, with
the aim of establishing a regional mechanism for research cooperation that supports the
sustainable development of capture-based aquaculture in the Asian region. Emphasis
has been placed on technology transfer and management strategies for the benefit of
farmers and coastal populations (Ottolenghi et al., 2004).
CONCLUSIONS
As a contributor to rural livelihoods, particularly those of coastal fishers, grouper
aquaculture can generate potentially large financial benefits. The high-value of grouper
on the export market ensures that farmers are able to generate a profit even when stocks
suffer heavy mortalities. Despite high initial investment costs, studies have shown that
with appropriate support, even the poorest can benefit from grouper culture, with
implications for both household well-being and community development. However,
based on the information reviewed in this report, capture-based aquaculture may
TABLE 21
Southeast Asia national regulations Locality Regulation
China
Limits the number of grouper “seed” fishers and the quantities of grouper
“seed” captured
A licence is needed for transporting marine “seeds” and their export is
prohibited
There is a management regulation of Guangdong Province for the cultivation
of aquatic products in the shallow sea intertidal zone, which applies to those
engaged in marine cultivation
China Hong Kong SAR Culturists must be licensed and operate in one of 26 gazetted culture zones
There are no regulations that apply to the capture of grouper “seeds” or their
import or export
Indonesia There is no management of seed resources
Malaysia Federal legislation prohibits the use of cyanide for fishing
In East Malaysia there are no special regulations for grouper seed capture.
Some regulations may act indirectly, for example some gears that are made
of trawl net are subject to trawl mesh size control. Grouper seeds cannot be
imported for culture
In West Malaysia the fishing of “seeds” is not allowed during November and
December; it is only permitted during the peak season from January to April.
No export of seeds smaller than 15  cm is permitted
Philippines It is illegal to use cyanide or any other poisonous substance for fishing
The scissor net is illegal
Fyke net have been banned
The Fisheries Code of 1998 (Republic Act 8550) prohibits the export of “seed”
of milkfish and prawn but its application to grouper is not clear. This Code
regulates gear/structures and operational zones for fish capture and culture
Transportation and export of fish and fisheries products requires permits from
the Quarantine section, including a health certificate from the Fish Health
section of Bureau of Fisheries and Aquatic Resources (BFAR)
Taiwan PC In Penghu Island, fishers are not permitted to catch any grouper seed of <6 cm
The use of cyanide for fishing is illegal
Thailand The use of push nets and fyke nets is limited. Push nets and trawlers should
not be used within 3 km of the shore and the mesh size of trawlers should be
≥2.5 cm.
Viet Nam Government regulations prohibit export of groupers <500  g (ex Ministry of
Fisheries)
There is no limit on export volumes. For export a health certificate from a
provincial office, Fisheries Resources and Environment Conservation Sub-
Department is needed, and requirements of the importing country satisfied
Source: Sadovy, 2000, as reprinted in Ottolenghi et al., 2004.

Capture-based aquaculture: global overview250
not be the best means to ensure a steady and sustainable supply of grouper for either
the live or “non-live” fish trades. This is due to a number of problems including low
availability of seed, destructive and wasteful seed collection techniques, removal of
large numbers of early life history stages with subsequent impacts on adult populations
and conflicts with capture fisheries, and pollution and disease resulting from culture
operations.
The obvious solution to some of the problems of CBA is to develop closed-cycle
hatchery rearing for all grouper species. Important advances in full-cycle culture have
been made for several species, particularly in Taiwan  Province of China, and full-cycle
culture appears financially feasible given a large enough capital investment. However,
given the financial means of most grouper culturists, and the difficulty in rearing most
grouper species, it remains unlikely that many of these species will be hatchery-reared
in the near future. It is also likely that hatchery production would undermine the
potential contribution of grouper culture in the livelihoods of the poor. Production
would most likely be taken out of the hands of small-scale producers. An increase
in production if hatchery fry is available would also lead to increased supply and a
likely drop in value, and lower profits. The market value of grouper is driven by its
relative rarity. On the other hand, poorer farmers would probably continue to fish for
grouper fry as they cannot afford to buy fry, and wild capture makes grouper culture
less risky and more accessible. In the meantime, steps must be taken to improve the
management of both CBA and capture fisheries for grouper. Some countries, such
as Palau, have taken strong measures to protect their reef fish populations, including
the closure of spawning seasons and spawning aggregation sites, bans on the export
of grouper and other vulnerable species, and even complete moratoria on fishing for
species in an obvious state of decline (e.g. humphead wrasse, Cheilinus undulates, and
bumphead parrotfish, Bolbometopon muricatum). This has effectively stopped the live
reef fish trade in Palau. In addition, the government of Palau, in cooperation with the
governments of the US and Japan, has developed viable full-cycle culture for at least
one commercially important grouper (Epinephelus fuscoguttatus), and experimentation
continues with other species.
Similar to export bans on adult grouper fisheries, Sadovy (2001b) suggested that all
export of grouper seed should be banned and that grouper should be cultured to market
size within their own country. This would allow for more stringent management of
grouper CBA, while reducing the transmission of disease via exported seed. Reduction
or elimination of the more wasteful and destructive seed collection techniques (e.g.  fyke
nets and scissor nets) is another appropriate step. Lastly, both CBA and capture
fisheries should promote the application of the precautionary principle and adopt the
FAO international Code of Conduct for Responsible Fisheries (CCRF).
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