217

Capture-based aquaculture of
groupers

Mark Tupper
WorldFish Center
Penang, Malaysia
E-mail: m.tupper@cgiar.org

Natasja Sheriff
WorldFish Center
Penang, Malaysia
E-mail: n.sheriff@cgiar.org

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
218 Capture-based aquaculture: global overview

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.
220 Capture-based aquaculture: global overview

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

Cromileptes altivelis (Valenciennes, 1828)

FIGURE 1
Humpback grouper (Cromileptes altivelis)

COURTESY OF J.E. Randall

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).

FIGURE 2
Distribution of Cromileptes altivelis (FishBase, 2007)
222 Capture-based aquaculture: global overview

Epinephelus akaara (Temminck and Schlegel, 1842)

FIGURE 3
Hong Kong grouper (Epinephelus akaara)

COURTESY OF A. Cornish
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).

FIGURE 4
Distribution of Epinephelus akaara (FishBase, 2007)
Capture-based aquaculture of groupers 223

Epinephelus amblycephalus (Bleeker 1857)

FIGURE 5
Banded grouper (Epinephelus amblycephalus)

COURTESY OF CSIRO
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).

FIGURE 6
Distribution of Epinephelus amblycephalus (FishBase, 2007)
224 Capture-based aquaculture: global overview

Epinephelus areolatus (Forsskål, 1775)

FIGURE 7
Areolate grouper (Epinephelus areolatus)

COURTESY OF J.E. Randall

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).

FIGURE 8
Distribution of of Epinephelus areolatus (FishBase, 2007)
Capture-based aquaculture of groupers 225

Epinephelus awoara (Temminck & Sclegel 1842)

FIGURE 9
Yellow grouper (Epinephelus awoara)

COURTESY OF D.C. Cook

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).

FIGURE 10
Distribution of Epinephelus awoara (FishBase, 2007)
226 Capture-based aquaculture: global overview

Epinephelus bleekeri (Temminck & Sclegel 1842)

FIGURE 11
Duskytail grouper (Epinephelus bleekeri)

COURTESY OF J.E. Randall

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).

FIGURE 12
Distribution of Epinephelus bleekeri (FishBase, 2007)
Capture-based aquaculture of groupers 227

Epinephelus bruneus (Bloch, 1793)

FIGURE 13
Longtooth grouper (Epinephelus bruneus)

FAO
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).

FIGURE 14
Distribution of Epinephelus bruneus (FishBase, 2007)
228 Capture-based aquaculture: global overview

Epinephelus coioides (Hamilton, 1822)

FIGURE 15
Orange-spotted grouper (Epinephelus coioides)

COURTESY OF D.C. Cook

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).

FIGURE 16
Distribution of Epinephelus coioides (FishBase, 2007)
Capture-based aquaculture of groupers 229

Epinephelus fuscoguttatus (Forsskål, 1775)

FIGURE 17
Brown-marbled grouper (Epinephelus fuscoguttatus)

COURTESY OF J.E. Randall

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).

FIGURE 18
Distribution of Epinephelus fuscoguttatus (FishBase, 2007)