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Sea cucumber culture, farming and sea ranching in the tropics: Progress,
problems and opportunities

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DOI: 10.1016/j.aquaculture.2012.08.053

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Aquaculture 368-369 (2012) 68–81

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Review

Sea cucumber culture, farming and sea ranching in the tropics: Progress, problems
and opportunities
Steven W. Purcell a,⁎, Cathy A. Hair b, David J. Mills c, d
a
National Marine Science Centre, Southern Cross University, PO Box 4321, Coffs Harbour NSW 2450, Australia
b
School of Marine and Tropical Biology, James Cook University, Townsville 4811 QLD, Australia
c
The WorldFish Center, PO Box 500 GPO, 10670 Penang, Malaysia
d
ARC Centre of Excellence for Coral Reef Studies, James Cook University, Townsville 4811 QLD, Australia

a r t i c l e i n f o a b s t r a c t

Article history: Tropical sea cucumber mariculture has potential to become a profitable industry and contribute towards natural
Received 8 March 2012 population replenishment. Here, we synthesise the fields of progress, current impediments and research oppor-
Received in revised form 31 August 2012 tunities in tropical sea cucumber aquaculture arising from recent studies and an Indo-Pacific symposium. We
Accepted 31 August 2012
present novel comparisons of data from hatcheries, earthen ponds and sea pens from published and unpublished
Available online 8 September 2012
studies in various countries. Of the few tropical species to have been cultured, only the sandfish Holothuria scabra
Keywords:
has been bred extensively. While risks from hatchery-produced sea cucumbers are recognised, more genetic re-
Holothurian search is needed in farming and sea-ranching programmes. Advances have been made in the culture and nursery
Co-culture rearing of tropical sea cucumber juveniles but few have been published. Sandfish larvae have now been grown
Pond culture successfully on just one microalga species, but experimental studies to optimise culture conditions are needed
Sea ranching urgently. Disease of tropical sea cucumbers in culture is infrequent but the treatment of disease and parasites
Mariculture is understudied. Earthen ponds are currently most effective for nursery rearing of juvenile sandfish to a size
Economic analysis for stocking. Growth rates and survival of sandfish in ponds to market size are also favourable, and should im-
prove via studies on stocking density, feeding regimes and pond management. Sea pens confer ownership of re-
leased stock and can provide a means of limiting predation in natural habitats but the costs of materials,
maintenance and surveillance against poaching can diminish profitability. Sea ranching has minimal material
costs but needs a large leased area and may require juveniles to be marked prior to release. Retail prices of
sandfish in Hong Kong increased exponentially with body size. A cost-benefit analysis illustrated that labour
and utility costs in pond farming will preclude profitability of monoculture in some cases, forcing proponents
to look towards co-culture or gamble with uncertain survival in sea ranching. Better governance and consultation
regarding the stocking of sea cucumbers have been advocated. We conclude that well-designed experiments and
meta-analyses are needed to fill critical knowledge gaps if sea cucumber mariculture is to expand in the tropics as
it has in temperate Asia. Co-culture remains a burgeoning frontier despite poor success of initial studies. Sea cu-
cumbers have superb potential to diversify mariculture industries in the tropics and potentially ameliorate the
detrimental effects of mariculture on coastal ecosystems.
© 2012 Elsevier B.V. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
2. Hatchery production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
2.1. Limitation of technical advances . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
2.2. Genetic risks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
2.3. Broodstock conditioning and spawning . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
2.4. Biological knowledge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
2.5. Larval and early-juvenile culture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
2.6. Diets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
2.7. Juvenile nursery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

⁎ Corresponding author. Tel.: +61 2 6648 3900; fax: +61 2 6651 6580.
E-mail address: steven.w.purcell@gmail.com (S.W. Purcell).

0044-8486/$ – see front matter © 2012 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.aquaculture.2012.08.053
 Author's personal copy

S.W. Purcell et al. / Aquaculture 368-369 (2012) 68–81 69

2.8. Disease and parasites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

3. Sea farming . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
3.1. Pond grow-out . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
3.2. Co-culture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
3.3. Farming in sea pens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
3.4. Sea ranching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
3.4.1. Genetic risks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
3.4.2. Other concerns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
3.4.3. Improving post-release survival . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
3.4.4. Prospects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
4. Sociological issues of stocking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
4.1. Governance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
4.2. Consultation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
4.3. Poaching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
5. Economics of sea cucumber mariculture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
5.1. Economic viability . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
5.2. Improving returns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
6. Conclusions and future opportunities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Appendix A. Supplementary data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

1. Introduction
While resource managers struggle to address pandemic
overfishing of tropical sea cucumber stocks (reviews in Purcell,
2010; Purcell et al., in press; Toral-Granda et al., 2008), mariculture
of some tropical species ascends as a viable prospect to subsidise
waning captures. Populations of high-value sea cucumbers have
been so decimated in some areas that better fisheries governance
and regulatory measures alone may be incapable of restoring
populations (Friedman et al., 2011; Hasan, 2005). Fortunately, tech-
nology for commercial-scale aquaculture has been proven for several
of the valuable, overfished tropical species (Agudo, 2006; Ivy and
Giraspy, 2006; Mercier et al., 2004) and culture methods are begin-
ning to develop for several other species (Table 1). Mariculture
may thus also provide a pathway to the rebuilding of wild stocks
via deliberate culture-based restocking or as a by-product of sea
ranching (Bell et al., 2005; Bell et al., 2008). Technology for culturing
a few of the tropical species has advanced in recent years but a num-
ber of diverse impediments currently undermine profitability
and reliability of mariculture and present challenging research
opportunities.
For centuries, sea cucumbers have been a popular luxury food item
in Asian dried seafood markets (Conand, 2001; Ferdouse, 2004;
Raison, 2008), where most of the product is traded and sold in the
dried form, called beche-de-mer or trepang. Of the few temperate spe-
cies in trade, the Japanese sea cucumber Apostichopus japonicus com-
mands superior prices up to about US$2950 kg −1 dried (Fig. 1a), but
prices vary (Purcell, unpubl. data). Consequently, research on culture
technology for A. japonicus has surged in China. Sophisticated Chinese
hatcheries now culture more than 6 billion juveniles annually (J. Chen,
pers. comm.) and production through sea ranching, cage culture and
pond farming is presently equivalent to one-quarter of global wild cap-
tures by dry weight (Purcell et al., in press).
More than 50 tropical species are commonly traded (Purcell, 2010)
but generally fetch relatively low prices, and smaller specimens command
lower prices per unit weight than larger ones. Notwithstanding, sandfish
Holothuria scabra is the most valuable and sells in Hong Kong for US
$115–640 kg−1 dried (Fig. 1b), and very occasionally as high as US
$1670 kg−1 for the largest specimens (for animals of 1.5–2 kg fresh
weight, equivalent to 75–100 g dried) (Purcell et al., 2012). Also of high
value, the golden sandfish H. lessoni can command US$240–790 kg−1,
and species within the ‘teatfish’ group (e.g. H. fuscogilva, H. nobilis, H.
whitmaei) can retail for US$130–270 kg−1 dried in Hong Kong (Fig. 1b)
(Purcell et al., 2012). Since the vast majority of tropical species are
exported to Asian markets and take years to attain large adult sizes
(Purcell et al., 2012), sea cucumber mariculture is expected to remain fo-
cussed on the high-value species.
In 2003, the United Nations Food and Agriculture Organization
(FAO) convened a workshop on ‘Advances in Sea Cucumber Aquacul-
ture and Management’ (ASCAM) in Dalian, China. ASCAM showcased
significant breakthroughs in hatchery practices, nutrition and disease,
much of which concerned the temperate species A. japonicus. Howev-
er, critical gaps in knowledge limited mariculture profitability, reli-
ability and diversification, especially for tropical species (Lovatelli et
al., 2004).
Sea cucumber aquaculture is a recurring priority in the development
plans of numerous Indo-Pacific countries (Jimmy et al., 2012; Mills et al.,
2012). Consequently, investors must know about hatchery technology,
risks and constraints. Compared to temperate species, few recent
developments in mariculture of tropical sea cucumbers have been pub-
lished in the primary literature. For example, in Aquaculture since 2007,
21 articles have been published on A. japonicus yet only two were pub-
lished on H. scabra.
In an attempt to bring recent progress to the fore, the Australian
Centre for International Agricultural Research (ACIAR) and the Secretar-
iat of the Pacific Community (SPC) convened the Asia-Pacific Tropical
Sea Cucumber Aquaculture Symposium in 2011. It marshalled more
than 20 experts in tropical sea cucumber aquaculture, stocking and
trade. The most recent developments in hatchery production technolo-
gy, release strategies, farming techniques, management practices, post-
harvest technologies for value adding, and supply chains and marketing
were presented. In addition to seminars and proceedings articles (Hair
et al., 2012), workgroup sessions allowed the 60 participants from 20
countries to identify knowledge gaps and constraints for future
research.
Past literature on the culture and stocking of tropical sea cucum-
bers has been reviewed elsewhere (Battaglene and Bell, 2004; Bell
et al., 2005; Raison, 2008). Here, we review advances in tropical sea
cucumber mariculture from recent publications and papers presented
at the ACIAR-SPC symposium. We also gathered recent published data
and unpublished data from mariculture programmes on H. scabra in
the Indo-Pacific (Table 1) to provide new comparisons of survival at
various stages in hatcheries, growth in earthen ponds, survival and
growth in sea pens, and analyses of profitability. The implications of
the recent advances and our analyses are discussed in the context of
research priorities arising from the symposium's workgroup sessions.
 Author's personal copy

70 S.W. Purcell et al. / Aquaculture 368-369 (2012) 68–81

Table 1
Estimated hatchery production of juvenile sea cucumbers globally, and approximate years of operation. n/a = not available.

Country Species cultured Annual production Use of juveniles Proponents Start year to
of 1 g juveniles end year

Australia (Northern Territory) Holothuria scabra 62,000+ Sea ranching; pond farming Tasmanian Seafoods Pty. Ltd. 2004–ongoing
Australia (Queensland) H. scabra 500,000 Sea ranching Bluefin Seafoods 2003–2009
Australia (Queensland) H. lessoni 330,000 Sea ranching Bluefin Seafoods 2004–2009
Australia (Queensland) H. scabra 1000 Experimental QLD DPI&F 2004–2007
Canada Parastichopus californicus n/a Pond farming Sustainable Ecological Aquaculture (SEA) 2009–ongoing
China Apostichopus japonicus >6 billion Sea ranching; pond farming Government and private hatcheries 1990–ongoing
Ecuador Isostichopus fuscus n/a Experimental n/a 2002–2008
Fiji H. scabra 500 Experimental Hunter Pearls, Fiji MAFF 2008–2010
FSM (Pohnpei) H. scabra 10,000 Experimental College of Micronesia, Land Grant Program 2009–ongoing
FSM (Yap) Actinopyga sp. n/a Stock enhancement n/a 2007
India (Tuticorn) H. scabra 3000 Experimental Central Marine Fisheries Research Institute 1988–2006
India (Tuticorn) H. spinifera na Experimental Central Marine Fisheries Research Institute 2001–2006
Iran (Bandar-e Lengeh) H. scabra na Experimental Persian Gulf Molluscs Research Station 2011
Japan A. japonicus >3 million Stock enhancement n/a 1977–ongoing
Kiribati H. fuscogilva 500–8000 Stock enhancement Kiribati Ministry of Fisheries 1997–2009
Madagascar H. scabra 200,000 Sea farming (pens) Blue Ventures, TMD, MH.SA 2007–ongoing
Maldives H. scabra 5 million Sea ranching Masmeeru Pty Ltd 1997–ongoing
Mexico I. fuscus 300,000 Pond farming Acuacultura Dos Mil 2008–ongoing
New Caledonia H. scabra 18,000 Experimental WorldFish Center 2000–2006
New Caledonia H. scabra 450,000+ Sea ranching; pond farming Société d'élevage aquacole de la Ouenghi 2011–ongoing
New Zealand Australostichopus mollis n/a Experimental National Institute of Water and Atmosphere 2007–ongoing
Palau Actinopyga mauritiana 500,000 Stock enhancement Government hatchery, Korean technicians 2009–2011
Palau Actinopyga miliaris 50,000 Stock enhancement Government hatchery, Korean technicians 2009–2011
Philippines (Bolinao) H. scabra 32,000 Sea ranching University of the Philippines MSI 2001–ongoing
Philippines (Mindanao) H. scabra 15,000 Sea ranching; pond farming University of the Philippines, DOST, PCARMD 2009–ongoing
Philippines (Bolinao) Stichopus horrens 500 Experimental University of the Philippines MSI 2009–ongoing
Philippines (Dagupan) H. scabra 20,000 Experimental NIFTDC-NFRDI 2009–2011
Philippines (Iloilo) H. scabra 11,000 Experimental SEAFDEC 2010–ongoing
Saudi Arabia H. scabra n/a Sea ranching National Prawn Company n/a
Solomon Islands H. scabra n/a Experimental WorldFish Center 1996–2000
USA (Alaska) P. californicus n/a Experimental Alutiiq Pride Shellfish Hatchery 2010–ongoing
Vietnam H. scabra 200,000+ Pond farming RIA3 2001–ongoing

2. Hatchery production
2.1. Limitation of technical advances
Sandfish, H. scabra, is the most commonly cultured tropical species
(Table 1) (Raison, 2008). The golden sandfish, Holothuria lessoni,
commands similarly superior retail prices (Fig. 1b) but few attempts
have been made to culture it (Ivy and Giraspy, 2006). White teatfish,
Holothuria fuscogilva, have been cultured in small numbers in Kiribati
since 1997 but the technology has been little documented (Friedman
and Tekanene, 2005). In Ecuador, Isostichopus fuscus has been cul-
tured but production is not at commercial scale (Mercier et al., 2004).
While much research has been published recently on methods for
upscaling the production of the Japanese sea cucumber, A. japonicus,
(e.g. Choi et al., 2009; Li and Li, 2010; Xia et al., 2012; Yuan et al.,
2010) technology for culturing tropical species has lagged. Studies in
the late 1990s on the sandfish in Solomon Islands developed methods
for culturing, detachment and grading, and early juvenile rearing
(Battaglene, 1999; Battaglene and Seymour, 1998; Battaglene et al.,
1999). However, few recent studies have progressed hatchery technol-
ogy of tropical species such as sandfish within an experimental research
approach.
spawned oocytes in contact with sperm from different males separately.
For sandfish, 30–50 broodstock are often used for spawning (Agudo,
2006; Duy, 2012) owing to practical limitations of hatcheries. Common-
ly, only a small proportion of those broodstock will be successfully in-
duced to spawn on any given attempt (Eeckhaut et al., 2012; Gamboa
et al., 2012; N. Agudo, unpubl. data; C. Hair, unpubl. data). Limited ge-
netic diversity of cultured sea cucumbers is likewise inherent if only a
few parents are stripped of gametes for artificial fertilisation (Eeckhaut
et al., 2012; Léonet et al., 2009). These problems are most concerning
where cultured animals can mix and breed with conspecifics in the wild.
Much of the literature on genetic risks of stocking hatchery-reared
marine animals concerns fish (Hindar et al., 1991; Le Vay et al., 2007).
Best-practice guidelines on how many sea cucumber broodstock to
use and how to manage spawning to avoid sperm dominance from
certain males are needed for hatcheries. Inevitably, selective breeding
(genetic improvement) of favoured strains of sea cucumbers will
arise, so guidelines should consider whether hatcheries have obliga-
tions about how that occurs. Genetic research needs to feature prom-
inently in future aquaculture and stocking programmes for sea
cucumbers for these concerns to be addressed.
2.3. Broodstock conditioning and spawning

2.2. Genetic risks
For programmes stocking marine animals in the sea, the number of
broodstock needed to avoid genetic alterations of wild populations
will vary among circumstances (Bell et al., 2005) but Le Vay et al.
(2007) suggest 50 broodstock as a general minimum, with equal num-
bers of males and females. Moreover, fertilisation of oocytes should be
controlled to maximise the breadth of parentage by placing portions of
Most commonly, sea cucumber broodstock are conditioned in ponds
or tanks prior to induced spawning in hatcheries or induced in tanks di-
rectly after being captured from the wild (Agudo, 2006; Morgan, 2000).
Broodstock conditioned in tanks can synchronise gametogenesis but
should be held at low density to avoid excessive weight loss (Duy,
2012). Induction of spawning in small tanks is achieved with a variety
of stressors including drying of broodstock, thermal shock, introduction
of algae powder, rapid changes in water depth in the tanks and spraying
 Author's personal copy

S.W. Purcell et al. / Aquaculture 368-369 (2012) 68–81 71

among tropical species, which have widely differing biological and

ecological traits (Conand, 1990; Purcell, 2010), and research compar-
ing technologies for a range of species is needed.

2.5. Larval and early-juvenile culture

Larval culture of tropical species is still hampered by poor survival

of planktonic larvae to the juvenile stage (Fig. 2). Rates of survival are
typically poor in the early larval stages and commonly around 1% to
the early juvenile stage, at which time they are normally transferred
from larval tanks to nursery raceways or hapas in ponds. The highest
reported rates of larval survival were from stripped females in Mada-
gascar. The wide variation in average survivorship among sandfish
programmes in different countries (Fig. 2) may reflect geographic
variation as much as hatchery practices.
The density of larvae in tanks is often between 0.3 and 1 ind ml −1
(Agudo, 2006) and trials indicate that survival to settlement relates
inversely to stocking density (Battaglene et al., 1999; Duy, 2012; W.
Bowman, pers. comm.). Replicated experiments are needed to under-
stand the relationships between larval densities in the planktonic
phases (i.e. blastula, gastrula, auricularia, doliolaria) and survival.
Density-dependent growth and survival may also vary among larval
stages. Experiments are also needed to compare survival of settled
pentactula larvae among different methods of transfer to nursery
tanks.
Methods for growing and feeding newly-settled sea cucumbers
have, for the most part, been adopted from methodologies for culturing
herbivorous gastropods like abalone (Battaglene, 1999). We question
whether benthic diatoms, as used for abalone, provide optimal growth
and survival for juvenile sea cucumbers. Indeed, little is known about
what dietary components are assimilated by small juvenile sea cucum-
bers. Watanabe et al. (2012a) showed that sandfish juveniles (~2 g)
grew better on shrimp tank detritus or shrimp faeces than a benthic
Fig. 1. Dried sea cucumbers for sale in retail shops in Hong Kong; a) Apostichopus japonicus,
b) Holothuria lessoni, H. scabra, H. whitmaei, H. fuscogilva and Isostichopus badionotus for sale 100
next to dried shark fin. Prices are in HK$ per 600 g (i.e. per pound). Photos: S. Purcell.

with water jets (Agudo, 2006; Duy, 2010). Despite extreme treatments,
broodstock do not always respond by spawning and, quite often, a rela-
tively small proportion of animals ejaculate or ovulate (Eeckhaut et al., 10
2012; N. Agudo unpubl. data). Alternatively, oocytes and sperm can be
Survival (%)

stripped from adults and gametogenesis induced artificially (Eeckhaut

et al., 2012; Léonet et al., 2009). While seasonal spawning of tropical
sea cucumbers is the norm (e.g. Battaglene et al., 2002; Conand, 1993;
Muthiga and Kawaka, 2009), at least I. fuscus in Ecuador is known to
spawn predictably throughout the year, making hatchery spawning 1
more reliable (Mercier et al., 2012).
Effective conditioning of sandfish broodstock can improve spawning
success (Gamboa et al., 2012). However, Morgan (2000) found that
broodstock fecundity and the hatch rate of fertilised eggs in sandfish
relate inversely to the period of captivity in tanks. Still lacking is a clear
0.1
understanding of conditions required for maintaining healthy broodstock
te ic g

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at hatcheries. More research is also needed on husbandry techniques to

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synchronise gametogenesis so that a large proportion of the animals

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spawn when required.

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2.4. Biological knowledge
In spite of two decades of breeding and research, the biological
and ecological requirements of tropical sea cucumbers in captivity re-
main understudied. For example, we know little about nutritional re-
quirements, feeding mechanics and food assimilation in the sandfish
larvae or juveniles. In addition to empirical studies describing these
aspects, practitioners will need to have the implications of such infor-
mation simplified in manuals. Hatchery requirements will also vary
Madagascar, 2011 - Eeckhaut, Lavitra and Pascal, unpubl. data
Mindanao, Philippines, 2009-2010 - Gamboa et al. (2012)
New Caledonia, 2002-2004 - N. Agudo, unpubl. data
Cairns, Australia, 2006-2007 - C. Hair, unpubl. data
Pohnpei, FSM, 2009-2011 - M. Ito, unpubl. data.
Madagascar, 2009-2010 - Eeckhaut et al. (2012)
, unpubl. data
Iloilo, Phillippines, 2010-2011 - J. Altamirano
Fig. 2. Survival of sandfish (H. scabra) in the hatchery at various stages of development from
different localities. Data sources given in legend.
 Author's personal copy
72 S.W. Purcell et al. / Aquaculture 368-369 (2012) 68–81
diatom (Navicula ramosissima). Further research should examine per-
formance and feeding of juveniles on various biofilms and feed types
and the role of bacteria in their diet. Sand added to tanks can also im-
prove the growth of juvenile sandfish (Watanabe et al., 2012a).
Hence, a variety of husbandry practices, such as water exchange re-
gimes and addition of sand, need to be compared experimentally.
2.6. Diets
Research on diets, nutrition and feeding of the auricularia stage in
tropical sea cucumber larvae is also critically lacking. Battaglene
(1999) suggested that up to four microalgae species could be fed to
larval sandfish at concentrations from 20,000 to 40,000 cells ml −1.
Correspondingly, a cocktail of Chaetoceros muelleri, Rhodomonas
salina and Isochrysis galbana is recommended in published manuals
for the culture of sandfish (Agudo, 2006; Duy, 2010). Mixed
microalgal diets have also been used for rearing golden sandfish H.
lessoni (Ivy and Giraspy, 2006) and the Central/South American I.
fuscus (Mercier et al., 2012). However, Knauer (2011) found that
sandfish larvae in Australia survived well on C. muelleri alone com-
pared to a mixed microalgal diet. In Vietnam and the Philippines,
sandfish larvae have also recently been reared successfully with just
one species of Chaetoceros (Duy, 2012; Gamboa et al., 2012). While
seeming a breakthrough, replicated trials are needed to see if simpli-
fied feeding with one alga species is universally applicable or has an
impact on larval performance compared to multi-species diets. Gen-
erally, experimental studies are also needed to better validate optimal
densities of microalgae at each larval stage for different sea cucumber
species.
Hair et al. (2011) achieved some success in feeding sandfish larvae
and early juveniles using a commercial paste mix of Isochrysis sp.
(30%), Tetraselmis sp. (20%), Pavlova sp. (20%) and Thalassiosira
weissflogii (30%). Algal pastes require refrigeration but will ultimately
reduce costs because culturing microalgae is a major limitation in
low-income countries and small island states. However, since the
paste microalgae settle out of the water column relatively quickly, alter-
native feeding strategies will be needed. Moreover, feeding experi-
ments will be needed since sandfish larvae may not assimilate some
paste microalgae such as Thalassiosira weissflogii (the replacement for
Chaetoceros in paste products) (J. Knauer, pers. comm.). Methods
developed by Watanabe et al. (2012b) to assess the condition of
juvenile sea cucumbers may assist in progressing this field of research.
2.7. Juvenile nursery
Nursery rearing of sea cucumbers in hatcheries to a size appropriate
for stocking in the sea is space-intensive; an issue which can be addressed,
for some species, through nursery culture in mesh enclosures in earthen
ponds. Early technology comprised two steps: small juveniles (3–5 mm
long) are firstly grown in fine-mesh ‘hapas’ (400–700 μm) then, at 1 g,
they are moved to larger-mesh (e.g. 1 mm) ‘bag nets’ (Pitt and Duy,
2004) and grown to 3–10 g for stocking to ponds or the sea. Fine-mesh
hapas can become clogged with fouling (Pitt and Duy, 2004), however,
with good cleaning practices, pond nursery in Vietnam is conducted sole-
ly in hapas (Duy, 2012). While the two-step method may minimise
cleaning requirements and escape of juveniles through the mesh, mate-
rials and labour for transfers are more expensive.
Within several concrete nursery tanks, Lavitra et al. (2010) found
that growth rates of sandfish juveniles were highest at lower stocking
density and relatively unaffected by sediment quality. The development
of nursery methods in earthen ponds has mostly been through
unreplicated trials. Sandfish juveniles should be greater than 5 mm in
length before transfer to 660-μm-mesh enclosures in ponds and upright
mesh structures within the enclosures can improve growth significantly
(S. Purcell and N. Agudo, unpubl. data). Experimental development of
cost-effective enclosures will be a key to scaling up commercial
production of juveniles. Alternative approaches, such as transferring ju-
veniles directly from larval rearing tanks to ponds via plastic sheeting
(Hair and Johnston, 2008), may prove more cost-efficient than current
methods but require further investigation.
The extreme variability in growth among juveniles (Gamboa et al.,
2012; Juinio-Meñez et al., 2012a; Pitt and Duy, 2004) is also problem-
atic for nursery culture. Successively removing the faster-growing ju-
veniles allows smaller individuals to grow more quickly (Battaglene
and Seymour, 1998), but grading techniques need to be optimised.
Feeding of juveniles in mesh enclosures also requires further testing
under different pond conditions. Fine shrimp feed may promote faster
growth of sandfish than other cheap feeds (Pitt and Duy, 2004; c.f.
Watanabe et al., 2012a). However, feeding may be unnecessary if
ponds have a rich natural productivity that provides organic detritus
in enclosures as food for sea cucumbers (S. Purcell and N. Agudo,
unpubl. data).
Where ponds are not available, mesh enclosures may be used in
the sea, however the technology is still nascent. Sandfish juveniles
reared in floating and bottom-set hapas at sea in the Philippines did
not survive or grow as well as in ponds (Juinio-Meñez et al., 2012a).
Similarly, slow growth and variable survival of sandfish juveniles in
mesh enclosures on the sea floor at New Caledonia indicated that fur-
ther experimentation with floating systems is needed (S. Purcell and
N. Agudo, unpubl. data).
2.8. Disease and parasites
Only relatively recently have diseases and parasites of cultured holo-
thurians been reported in the literature (Eeckhaut et al., 2004; Jangoux,
1990; Mercier et al., 2004; Wang et al., 2004). The ACIAR-SPC sympo-
sium underscored a number of serious health issues facing hatcheries
for tropical sea cucumbers:
(1) Outbreaks of the larvae of dipteran flies, colloquially called
‘bloodworms’, can compete with juvenile sea cucumbers for re-
sources in tanks (Gamboa et al., 2012). Abatement of dipteran
larvae by covering tanks with fine mesh has worked in some in-
stances but tends to be incomplete.
(2) Protozoans can parasitise juvenile sea cucumbers internally,
causing disease (Mercier et al., 2004). In South America, they ap-
pear to be limited by aeration and the control of water temper-
atures (Mercier et al., 2012).
(3) Bacterial infections on juvenile and adult sea cucumbers cause
skin lesions and eventual breakdown of the body wall (Agudo,
2012; Eeckhaut et al., 2004; Hair and Johnston, 2008; Purcell
and Eeckhaut, 2005). A body-wall disease in cultured I. fuscus in
Mexico and Ecuador was partly treatable with antibiotics but
the best prevention and cure was transfer to clean, oceanic
water (Mercier et al., 2012). However, potential effects on wild
populations should be investigated as body-wall diseases can be
highly contagious (Eeckhaut et al., 2004). In Australia and New
Caledonia, diseased sandfish were quarantined and successfully
treated by an antibiotic bath (e.g. oxytetracycline at 100 ppm
for 1 h) and by improving water quality (Morgan, 2000; C. Hair,
pers. obs., N. Agudo pers. comm.). Similarly, Eeckhaut et al.
(2004) recommend quarantining infected sandfish and a weak
formalin bath.
(4) Copepods compete for food in tanks with sea cucumbers. This
generic mariculture problem can be avoided by conditioning set-
tlement plates (prior to settlement of sea cucumber larvae) in a
sterile algal monoculture or by painting plates with a slurry of
commercially available Spirulina rather than natural diatoms
(Agudo, 2006; Pitt and Duy, 2004). Copepods can be treated
with brief doses of Trichlorfon/Dipterex at concentrations of
about 1–2 ppm (Agudo, 2006; James, 2004; Pitt and Duy, 2004).
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S.W. Purcell et al. / Aquaculture 368-369 (2012) 68–81
Chlorination–dechlorination treatment of seawater, filtration of
seawater to 1 μm, and manipulations of salinity and/or temperature
of stock seawater may also abate parasites and pests. Maintaining
good water quality and avoiding high stocking densities and handling
of juvenile sea cucumbers should help to avoid disease (Purcell and
Eeckhaut, 2005; Raison, 2008). Currently lacking in literature on
mariculture of sea cucumbers are case studies in which the control
of parasites and disease has been tested under different treatments,
e.g. chemical types, concentrations and durations.
3. Sea farming
3.1. Pond grow-out
Survival of cultured sandfish juveniles to an adult size tends to be
much higher in ponds than in the sea — usually around 50–85%
(Agudo, 2012; Bell et al., 2007; Duy, 2012; Gamboa et al., 2012).
The sandfish is amenable to pond conditions as it is an inshore species
of silty habitats (Agudo, 2012; Hamel et al., 2001; Raison, 2008). In
contrast, most other commercially valuable tropical species inhabit
coral reef environs (Conand, 1990) and will likely not tolerate
muddy pond substrates or pond seawater. For example, Pacific black
teatfish (Holothuria whitmaei) broodstock, averaging 1.8 kg body
weight, lost on average 180 g each over 6 months in the same earth-
en pond in which sandfish gained weight (N. Agudo, unpubl. data).
Similarly, white teatfish (H. fuscogilva) do not survive or grow well
in milkfish ponds in Kiribati (K. Aram, pers. comm.). Experiences in
Ecuador and Mexico (Mercier et al., 2012) suggest that earthen
ponds offer mediocre conditions for grow-out of I. fuscus.
Sea cucumbers consume particulate organic waste matter in ponds
and some species (e.g. H. scabra) bioturbate pond sediments (Fig. 3;
Purcell, 2004a). Surprisingly, the assimilation of pond-sediment constit-
uents by sea cucumbers has been understudied. Sandfish in earthen
ponds seem to efficiently extract nutrients from enriched sediments
and grow relatively fast. For example, in New Caledonia, growth of juve-
nile and small adult sandfish was 20–72 g month−1 in ponds (Agudo,
2012; Bell et al., 2007; Purcell, 2004a) but about 14–22 g month−1
over 12 months in coastal seagrass beds (Purcell and Simutoga, 2008).
Comparison of most recent data shows strong correspondence
of growth rates of sandfish in earthen ponds among trials in New
Caledonia, Vietnam and Australia (Fig. 4). Absolute growth rates
of small juveniles (i.e. 5–50 g) tend to be slow in the first few
months in ponds (e.g., 0.1–0.9 g ind − 1 day − 1) then increase
Fig. 3. Underwater photograph of an adult sandfish (H. scabra) half buried into upper sedi-
ments in an earthen aquaculture pond. The bioturbation of dark anoxic sediments through
burying action is evident. Photo: S. Purcell.
73
450
400
350
Mean body weight (g)
300
250
200
150
100
50
0
0 2 4 6 8 10 12 14 16 18
Grow-out period in ponds (months)
Vietnam - Pitt and Duy (2004)
New Caledonia - Bell et al. (2007)
Ayr, Australia - C. Hair, unpubl. data
India - James (1999)
Vietnam - N. Duy, unpubl. data
Mindanao, Philippines - Gamboa et al. (2012)
Fig. 4. Grow-out of cultured juveniles in earthen ponds. Average body weights (symbols)
and growth trajectories of cultured sandfish (H. scabra) in earthen ponds from different local-
ities. Data sources given in legend.
(e.g., 1.0–1.7 g ind − 1 day − 1) once they reach a size at which they
can ingest larger quantities of food (Agudo, 2012; Pitt and Duy,
2004; Purcell and Kirby, 2006; Raison, 2008). Seasonal declines in
seawater temperatures in ponds are a problem for growth of
sandfish at high latitude (Agudo, 2012; Bell et al., 2007) but not
at low latitude (Pitt and Duy, 2004) (Fig. 4). Growth rates of
sandfish in pond trials in Mindanao, the Philippines (Gamboa et
al., 2012), appear to be comparatively fast (Fig. 4), which may re-
flect consistently warm conditions. Proliferation of filamentous
benthic algae in ponds can retard growth or kill sandfish (Agudo,
2012; Hair and Johnston, 2008) and may be prevented by stimulat-
ing a microalgal bloom in the water column. Future research to im-
prove growth rates in ponds should, in particular, examine the
effects of stocking densities, water exchange and feeding.
Experimental studies in Vietnam (Pitt and Duy, 2004) were a cata-
lyst for commercial mono-culture of sandfish in rotational cycles with
shrimp, currently yielding 2–3 tonnes ha−1 y−1 (Duy, 2012). Howev-
er, stratification of water from heavy rains is a key problem for farmers.
Various methods have been trialled to ameliorate water stratification in
ponds with sandfish (Agudo, 2006; Pitt and Duy, 2004). Mills et al.
(2012) suggest regular tidal exchange and paddlewheels to de-stratify
pond water in Vietnam, although Duy (2012) claims that mixing from
tidal water exchange and wind can be sufficient. In northern Australia,
Bowman (2012) used air diffusers and drained freshwater from the sur-
face of ponds. Since interventions such as paddlewheels, air diffusers
and seawater pumping incur costs, replicated trials to compare cost-
effectiveness of water-stratification management strategies would be
widely useful.
3.2. Co-culture
Co-culture is attractive over monoculture of sea cucumbers owing
to their requirement for relatively low density culture (Battaglene et
al., 1999), a relatively long culture cycle to a large (higher price per
unit weight) market size, their sedentary benthic habit, their ability
to feed on faeces and pseudofaeces (Raison, 2008; Slater and Carton,
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74 S.W. Purcell et al. / Aquaculture 368-369 (2012) 68–81
2007) and apparently low rates of emigration from co-culture sites
(Slater and Carton, 2010). Technology for culturing sea cucumbers
under, or with, other taxa has advanced considerably for temperate
species (e.g. Paltzat et al., 2008; Zamora and Jeffs, 2011; Zhou et al.,
2006). Unfortunately, similar technology for tropical species has
failed to develop.
Co-culture trials of sandfish with Babylon snail Babylonia areolata in
Vietnam initially yielded poor survival (Pitt and Duy, 2004). Also in
Vietnam, juvenile sandfish did not survive well with tiger prawns
Penaeus monodon (Pitt et al., 2004) nor with P. vannamei post-larvae
once shrimp attained a size at which they were capable of damaging
sandfish (Mills et al., 2012). In New Caledonia, sandfish juveniles
could co-exist with post-larval blue shrimp Litopenaeus stylirostris
(Purcell et al., 2006a) but not larger shrimp — neither dispersed within
ponds nor within mesh enclosures in ponds (Bell et al., 2007). Perhaps
the lack of success can be attributed to initiating co-culture too early
in the life cycle of sandfish, but it seems in hindsight that shrimp present
an unreliable partner for sea cucumber co-culture.
Studies on alternative options for culturing tropical sea cucumbers,
such as the sandfish, in co-culture with fish, shellfish or algae should
be supported. Mills et al. (2012) reported that sandfish grew successfully
in pond trials with sea bass (Lates calcarifer or barramundi) in Vietnam.
Likewise, preliminary co-culture with milkfish (Chanos chanos) and
pompano (Trachinotus blochii) showed promise, but not rabbitfish
(Siganus guttatus) or grouper (Epinephelus coioides). Future co-culture
research should test different stocking densities over a range of animal
sizes of both co-cultured species. Moreover, co-culture of tropical sea cu-
cumbers may not prove to be a panacea, since it is conceivable that more
soluble wastes are liberated from farms with sea cucumbers (Purcell,
2004a) or that stocking densities may need to be reduced in co-culture.
Hence, cost-benefit analyses will be required as evidence of economic
advantage.
Issues of managing ponds or cage mariculture with species having
faster culture cycles than sea cucumbers also need to be considered. In
Vietnam, grow-out of sandfish to the minimum market size in rotation-
al culture with shrimp in ponds receives a “head start” (Duy, 2012; Mills
et al., 2012) by firstly rearing juveniles to 50 g in nursery ponds so that
they require a relatively short grow-out period of 7–9 months once sold
to farmers. While rotational culture in ponds presents a profitable op-
portunity in some cases (Raison, 2008), the culture of tropical sea cu-
cumbers under the suspended culture of other invertebrates remains
a promising arena for development because they can be grown over
longer periods to sizes that command premium market prices.
3.3. Farming in sea pens
Sea pens retain sea cucumbers in a defined area while allowing tidal,
or current-driven, seawater exchange and access to naturally occurring
sediments, seagrass and organic detritus. In most cases, there is no
cover over the pens and, particularly if the walls are low, the sea pens
allow access to predators. Stiff plastic mesh has worked best and the
bottom of the pen mesh is often buried into the sediments to prevent
sea cucumbers from burying underneath and escaping (Purcell and
Simutoga, 2008; Robinson and Pascal, 2012). Three common applica-
tions of sea pens for tropical sea cucumber mariculture are as follows:
1) Grow-out of wild-captured juveniles. The pens constrain migration
of the sea cucumbers, designate ownership of stocks and deter
poachers. Large, high-fenced pens for this purpose have been
used in India (James, 1996), the Philippines and Indonesia (Bell
et al., 2008; Tuwo, 2004). The benefit of increased egg supply to
improve recruitment in the fishery can occur if fishers allow the
animals to grow beyond the size-at-maturity (Bell et al., 2008).
2) Grow-out of hatchery-produced juveniles. Sea pens for this purpose
have tended to be relatively large (156–900 m 2) and low (e.g. 1 m
high and buried 25 cm into sediments), and simply contain the
sea cucumbers but fail to exclude predators such as crabs
(Lavitra et al., 2009; Robinson and Pascal, 2009, 2012). The pens
may belong to individuals or family groups within a community
and may share adjoining walls.
3) A research tool for better estimating survival of juveniles. Smaller sea
pens (e.g. 1–500 m 2) can be used in the short period after releas-
ing hatchery-produced juveniles in the wild (Purcell, 2004b,
2012). The natural habitat is preserved inside the pens and they
are designed to retain juveniles but not affect other processes
such as predation, water quality fluctuations and food availability
(Purcell and Simutoga, 2008). Small, replicated experimental pens
can be used as pilot units (Purcell and Simutoga, 2008; Robinson
and Pascal, 2009) or pens of 100–500 m 2 for large-scale experi-
ments (Hair, 2012; Juinio-Meñez et al., 2012b; Purcell and
Simutoga, 2008).
Growth of sandfish in sea pens tends to be somewhat slower than in
earthen aquaculture ponds, presumably because natural sediments are
not enriched by leftover feeds and faeces, and there are more competi-
tors and predators. The fastest (albeit variable) growth rates of sandfish
in sea pens were reported from Madagascar and the slowest from New
Caledonia (Fig. 5a). In an exceptional case at low stocking density in
Madagascar, sandfish released at 15 g grew to an average of 420 g in
7 months (Robinson and Pascal, 2012). More commonly, sandfish
a
400
300
Body weight (g)
200
100
0
b 100
80
Survival (%)
60
40
20
0
0 2 4 6 8 10 12
Time from release (months)
New Caledonia - Purcell and Simutoga (2008)
Vietnam - Pitt and Duy (2004)
Madagascar - Robinson and Pascal (2012)
Fiji - Hair (2012)
Fig. 5. Grow-out of cultured juveniles in sea pens. (a) Average body weight (dots) and
growth trajectories (lines) and (b) survival of sandfish (H. scabra) released at various sizes
in sea pens in Vietnam, New Caledonia, Madagascar and Fiji. Data sources given in legend.
 Author's personal copy
S.W. Purcell et al. / Aquaculture 368-369 (2012) 68–81
juveniles of 1–10 g attain 150–250 g at 12 months after release
(Fig. 5a). In some localities, sandfish grew slower when seawater tem-
peratures cooled (Purcell and Simutoga, 2008) and, at other localities,
grew slower in hot conditions of summer (Hair, 2012). Some locations
may also not confer sufficient growth of sandfish to enable them to
reach market size in a reasonable timeframe (G. Robinson, pers.
comm.). The carrying capacity of sandfish in sea pens, in terms of bio-
mass of sea cucumbers per unit area, is difficult to define because it
will vary seasonally and among sites. Nonetheless, it has been reported
in the range of 200–250 g m−2 in New Caledonia (Purcell and
Simutoga, 2008) and 100–771 g m−2 at sites in Madagascar (Lavitra
et al., 2010; Robinson and Pascal, 2012).
Survival of sandfish in sea pens has also been highly variable but
most often poor in the first months after release (Fig. 5b) when the ju-
veniles are still small and perhaps behaviourally inept. Trials resulting
in high mortality rates tend to be reported with less detail than those
that achieved low mortality (e.g. Pitt and Duy, 2004). In Madagascar,
survival of sandfish in open sea pens averaged 48% over 2–11 months,
excluding those where sandfish were stolen (see Robinson and Pascal,
2012). In Fiji, sandfish in community-managed sea pens suffered 72%
mortality over the first 6 months (Hair, 2012). Projection of survival
trajectories in Fig. 5b indicates that survival of sandfish juveniles at
12 months after release will seldom be greater than 30%.
The major drawback of sea pens in comparison to sea ranching in
open habitats is the expense and effort required to construct, clean
and repair them (Robinson and Pascal, 2009). As with farming and sea
ranching, surveillance to prevent poaching is another considerable
cost. Sea pens are vulnerable to storms and rough seas (Hair, 2012;
Robinson and Pascal, 2012) and infrequent maintenance reduces their
lifespan. While the conditions within sea pens are similar to outside
the pen, providing a proxy for survivorship in the sea ranching situation,
this may not always be the case. Purcell and Simutoga (2008) speculate
that some predators may actually be attracted to sea pens. Moreover,
poorly positioned pens will disadvantage animals and prevent them
from migrating to better microhabitats.
3.4. Sea ranching
3.4.1. Genetic risks
Long-established alleles of local strains may confer adaptations to
local conditions or to infrequent environmental or biotic stressors
(Purcell, 2012). Losing those adaptations by out-breeding of local al-
leles from the intermixing and breeding of stocked animals of foreign
origin is a risk. Additionally, translocated marine animals released
into habitats in foreign waters may temporarily out-compete endem-
ic wild conspecifics (Hindar et al., 1991). Either mode of genetic alter-
ation could result in populations lacking the adaptations to cope with
infrequent local stressors. Hatchery-produced sea cucumber should
therefore only be stocked at sites with wild populations of similar ge-
netic structure (Purcell, 2012; Uthicke and Purcell, 2004).
Large genetic differences have been shown to exist among stocks
from different countries (Uthicke and Benzie, 2001; Uthicke and
Purcell, 2004) so inter-country translocations are strongly discouraged.
Nonetheless, there are a few instances where cultured juveniles from
parentage of one country's stock have been translocated to another coun-
try and released into the wild for sea ranching — and potentially negative
genetic impacts were exacerbated through deliberate fishing-down of
wild stocks (Jimmy et al., 2012; Pakoa et al., 2012). Countries and states
are urged to minimise harmful effects of introducing non-native species
and promote steps to minimise adverse genetic effects of cultured ani-
mals on wild stocks (FAO, 1995).
Concerns about genetic pollution due to inter-country transloca-
tions do not apply to cases where a species of sea cucumber may be in-
troduced to countries beyond its geographic distribution (Purcell,
2012). Indeed, translocations of other invertebrates, such as the gastro-
pod Trochus niloticus, throughout the Pacific have given rise to fisheries
75
that have proved important for coastal incomes. Likewise, the risk of
tropical sea cucumbers being ‘invasive’ species or ecological pests is
small because their populations attract intense fishing pressure and
the animals contribute positively to ecosystem functions (Purcell et
al., in press). If countries wish to import and introduce a new species
(e.g. H. scabra or H. lessoni) to local marine areas, the key questions are:
what is the nearest source population from which to procure broodstock?
And, what biosecurity protocols should be applied to minimise risk to the
other native sea cucumber species and marine biota?
3.4.2. Other concerns
Tank experiments indicate that sea cucumbers are beneficial to sed-
iment health (Uthicke, 1999). Additionally, field studies show that they
can improve seagrass productivity in some cases (Wolkenhauer et al.,
2010) and recycle nutrients for use by reef primary producers
(Uthicke and Klumpp, 1998). Their processing of sediments may also
contribute to the balance of CaCO3 on coral reefs, thereby reducing the
effects of ocean acidification (Schneider et al., 2011). While the ecosys-
tem benefits of aquaculture-based stocking are commonly touted, the
risks of stocking sea cucumbers in the wild have been rarely discussed
(Eriksson, 2012; Eriksson et al., 2012). Unforeseen impacts could include
shifts in benthic community structure from over-stocking, and introduc-
tion of disease, parasites or other foreign macro-associates. Importing
agencies should at least take actions to minimise the introduction of dis-
ease and parasites from cultured animals to wild populations (FAO,
1995). Surrogate protocols for best practice in translocating, quarantin-
ing and stocking can be used until these can be specifically developed for
sea cucumbers (Eriksson et al., 2012).
3.4.3. Improving post-release survival
Studies on releases of sandfish show that most cultured juveniles
do not survive to market size in the wild and that survival is variable
from one site to another, and potentially from one time to another
(Juinio-Meñez et al., 2012b; Purcell and Simutoga, 2008). In contrast
to research on hatchery methods, studies on ways to improve survi-
vorship of sea cucumbers in the sea are few indeed (Purcell, 2004b).
Sociological factors in community-based programmes are primary
criteria for site selection (discussed later). The habitat in which the
animals are released, the size of the animals, conditions of transport
and acclimation at release sites also appear to be critical (Pitt and
Duy, 2004; Purcell, 2012; Purcell and Simutoga, 2008; Purcell et al.,
2006b; Robinson and Pascal, 2012).
Research on behavioural conditioning of juveniles in the hatchery
and acclimation of the juveniles at sites prior to release could also
help to improve post-release survival. Juinio-Meñez et al. (2012a)
found that juveniles that had been conditioned on sand in the hatchery
buried more frequently in the first hour after release in the wild than
unconditioned juveniles. Such conditioning could help to instil natural
predator-avoidance behaviours and the effects on stocking success
could be tested; e.g. by estimating survivorship of fluorochrome-
marked controls and conditioned juveniles in open sea pens.
Predation of juvenile sea cucumbers by carnivorous fishes, birds, tur-
tles, sea stars, crabs, gastropods and other invertebrates (Dance et al.,
2003; Francour, 1997; Lavitra et al., 2009; Purcell and Simutoga, 2008)
is a principal disadvantage of sea ranching or sea-pen farming over
land-based farming in ponds. Site-specific knowledge of the key preda-
tors of cultured sea cucumbers could enable mitigation strategies. For ex-
ample, crabs may be able to be removed selectively (Robinson and Pascal,
2012), or refuges could be provided for the juveniles or they could be re-
leased at times of the year when predation risk is expected to be lower.
3.4.4. Prospects
As overfishing continues to diminish stocks of high-value sea cu-
cumbers in the tropics (Anderson et al., 2010; Purcell et al., in press)
and places more species in danger of extinction (Polidoro et al., 2011),
resource managers may need to close fishing of wild stocks. Sea
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76 S.W. Purcell et al. / Aquaculture 368-369 (2012) 68–81
ranching then faces a challenge of proving that harvests are from cul-
tured animals or risk opening opportunities for black marketing of
protected wild individuals and undermining conservation efforts
(Eriksson et al., 2012). Proof of ownership is likewise an issue where
sea ranching proponents need to pay resource owners for harvests of
wild stock. Marking the ossicles of juveniles by immersion in fluoro-
chrome solutions and subsequent laboratory analyses of tissue samples
provide a means to later distinguish the animals from conspecifics in
the wild (Purcell, 2012; Purcell et al., 2006c), but may have limited ef-
fectiveness after years (Purcell et al., 2006c). Scope remains for the de-
velopment of alternative marking or branding methods, especially for
application where proponents must identify hatchery-produced ani-
mals on sight underwater to be entitled to harvest.
Carrying capacity of sea cucumbers is a concern for resource man-
agers, especially where wild populations are still abundant. In the
Northern Territory, Australia, there are still good stocks of wild sandfish
amidst sea ranching projects (Bowman, 2012; Fleming, 2012), and
there are other localities where stock enhancement will be the aim
(e.g. Fiji, see Hair, 2012). A better understanding of the carrying capacity
of sea cucumbers in the wild will probably come from meta-analysis of
case studies of various stocking programmes and the bio-physical con-
ditions at a range of sites.
4. Sociological issues of stocking
4.1. Governance
Experience has shown that the social components of governance are
as critical as biological issues in establishing sustainable sea cucumber
farming (Eriksson et al., 2012; Juinio-Meñez et al., 2012b; Robinson
and Pascal, 2009). In this context, governance deals with the people
who are affected and empowering them to manage the grow-out and
harvest phases so that they can reap the profits (Eriksson et al., 2012;
Juinio-Meñez et al., 2012b). Coastal communities and villages engaged
in farming and sea ranching must have cohesion and effective internal
governance. A new frontier for this emerging sector is the development
of models for access rights and social equity.
Governance systems must provide adequate property rights and in-
centives to participants, balanced carefully against any negative and ex-
clusionary effects on non-participants. Aquaculture in any guise,
particularly in developing countries, has the potential to increase ineq-
uities by denying a section of the community access to a productive
asset (Luttrell, 2006) — this may be coastal land or ecosystem services
in the case of pond culture, or fishing grounds in the case of sea
ranching. Such inequities spur conflict and a lack of respect for regula-
tions, jeopardising sustainability.
Governance issues associated with the use of existing coastal ponds
are straightforward; ownership and leasing arrangements define clear
use rights. Governance of sea ranching is more complex because the
strategies must be context specific; integration with existing governance
institutions provides a robust, locally adapted ‘shortcut’ to this end. In
Fiji, customary tenure systems (tabu) implemented through ‘locally
managed marine areas’ (LMMAs) proved pivotal in establishing a reserve
to protect juvenile sea cucumbers released for ranching trials (Meo,
2012). In the Philippines, the provision of tenure for pilot ranches was
tied closely to existing institutions in the form of People Organisations,
Barangay (village) Councils and Local Government Units (Juinio-Meñez
et al., 2012b).
Partnership with industry is emerging as a mechanism to overcome
high capital requirements for establishing hatcheries while sharing
benefits with communities (Bowman, 2012; Fleming, 2012; Robinson
and Pascal, 2012). In Madagascar, the risk associated with sandfish mor-
tality and theft was entirely borne by those least able to absorb it — the
households (Eriksson, 2012). Negotiating equitable benefit and risk
sharing in such circumstances is critical, and may require external advo-
cacy on the part of communities. In northern Australia, governance
challenges of sea cucumber ranching include fostering equitable part-
nerships between Indigenous communities and industry, building ca-
pacity for communities to engage in the mainstream economy, and
mechanisms for ensuring that the vision and direction of the industry
are in the hands of communities (Fleming, 2012). Lastly, the need for
governance systems for sea cucumber farming to be integrated with
strong fishery and trade regulations, rather than being seen as a ‘fix’
or replacement for depleted fisheries, cannot be emphasised too strong-
ly (Eriksson, 2012; Juinio-Meñez et al., 2012b).
4.2. Consultation
Identifying and engaging with the stakeholders are critical initial
steps in establishing sea ranching or sea farming programmes. Especial-
ly for sea ranching, consultation needs to include fishers and managers
of the wild fishery and, as highlighted above, the stocking programme
integrated within the fishery management strategy (see Fleming,
2012; Hair, 2012; Juinio-Meñez et al., 2012b). Pascal and Robinson
(2011) recommend to pre-screen and select farmers for sea pen mari-
culture that can be expected to be dedicated, entrepreneurial types
committed to methodical work and regular expenses, and who accept
that income may be sporadic. Thereafter, performance-based criteria
after several cycles of farming could help to show which individuals
should remain involved in the activity (Pascal and Robinson, 2011).
Community-level education appears to be critical in providing an un-
derstanding of governance systems, an appreciation for shared benefits,
and adoption by communities to minimise conflict. In the Philippines,
education about the sea ranching extended beyond focus communities
to adjacent villages, allowing them to understand the potential benefits
of egg supply from the stocked adults in the sea ranch to the fishery
(Juinio-Meñez et al., 2012b). In Madagascar, local non-government or-
ganisations have produced a manual to guide communities in sea cu-
cumber farming (Pascal and Robinson, 2011). In Australia, education
initiatives in Indigenous communities participating in sandfish sea
ranching trials have extended to local primary schools to raise aware-
ness among the youth (Fleming, 2012).
Proponents of sea ranching and farming tend to be overly optimistic
about the likely returns to communities in whom they develop partner-
ships. Expectations of growth and survival of stocked sea cucumbers
must be realistic (Purcell, 2012) and balanced with discussion of the
wide range of risks and costs (Eriksson, 2012). Ideally, investors should
take on these risks, given their greater ability to recover from external
factors, and possibly offer a business model in which communities are
employed (Eriksson et al., 2012). The development and provision of
expert technical advice to sea cucumber farmers should help to reduce
risks involved with this fledgling industry.
4.3. Poaching
Theft of sea cucumbers from ponds, sea pens or sea ranches will be
common in low-income countries where people in neighbouring vil-
lages are financially desperate or jealous. Therefore, identifying who is
poaching, and understanding why, is important in forming communica-
tion strategies and risk mitigation measures (e.g. see Robinson and
Pascal, 2012). At least in wild sea cucumber fisheries, it is now apparent
that poverty is a key driver for the over-exploitation of stocks and man-
agement plans should be embedded in overarching aid aimed at pover-
ty reduction (Eriksson et al., 2012; Purcell et al., in press). Likewise, sea
ranching will face social conflict and poaching unless poverty and liveli-
hoods in neighbouring villages can be improved broadly. This situation
may be slightly better in some Pacific island situations where local ma-
rine tenure is still active and respected (Meo, 2012; Pakoa et al., 2012).
Despite good consultation, surveillance of stocked sea cucumbers
and enforcement of access rights will mostly be necessary. Australia
is one exception, where licensing controls and surveillance effectively
prevent poaching (Fleming, 2012). In low-income countries, such as
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S.W. Purcell et al. / Aquaculture 368-369 (2012) 68–81 77

Madagascar and the Philippines, surveillance of sea ranching or sea
pens may be handled at the community level (Juinio-Meñez et al.,
2012b; Robinson and Pascal, 2009). Such surveillance has required
villagers to guard released stock at night (e.g. Madagascar) or around
the clock (e.g. the Philippines), which can incur a cost. Surveillance
costs must be included at the project planning and evaluation stage
(Pascal and Robinson, 2011) and profit-sharing commensurate with ef-
fort in this role (see Juinio-Meñez et al., 2012b). In Madagascar,
poaching of sea cucumbers from coastal sea pens occurred after a natu-
ral disaster (cyclone) and during festivities, when surveillance was low.
Sadly, poaching may be intractably difficult to solve in some cases.
a
2.5
Vietnam
(US$ kg-1 production)
5. Economics of sea cucumber mariculture
5.1. Economic viability
The economics of growing sandfish in sea pens in Madagascar has
been briefly described by Pascal and Robinson (2011). To illustrate the
economics of sea cucumber production in ponds, we compared
bio-economic model outputs for similar hypothetical 8-ha farms in
New Caledonia and the central coast of Vietnam (Fig. 6). The models
were generated using decision-support software for prospective sea cu-
cumber farmers (Johnston, 2012), and employ a discounted cash-flow
approach to predicting returns over a specified enterprise lifecycle.
The farm structures reflect realistic parameters for enterprises based
on actual costs in both locations (Appendix S1). Briefly, the following
scenarios were compared:
Vietnam. Ponds were leased, more workers were needed but la-

2.0
bour costs were relatively low, and water exchange in ponds
1.5 was by tidal action. Prices for whole fresh sandfish varied by size
Net benefit

1.0 (VND52,000 (US$2.50) for 150–250 g; VND83,300 (US$4.00) for

250–400 g) with the market accepting relatively small-sized
0.5
sandfish. A 10-month growing period was practiced by farmers
0
to avoid the risk of low pond salinities in the wet season.
-0.5
-1.0 New Caledonia. Pond farms were part of an existing large-scale
180 230 280 330 380 aquaculture facility, managed under existing structures, and few,
Weight attained at 10 months (g) higher paid, staff were employed. Leasing costs were included in
calculations, and pond water was pumped from the sea into
Labour ponds using existing farm infrastructure. Accordingly, no capital
Juveniles
expenses are included in this model. Growth rates (see Agudo,
Operating
F.O.R.M. 2012; Bell et al., 2007) dictated an 18-month grow-out period to
Capital obtain a size acceptable for local buyers (500 g) and we used the
Electricity upper current price of 300 XPF (US$3.34) for whole fresh sandfish.
0 0.2 0.4 0.6 0.8 1 1.2 1.4 The model outputs demonstrate a stark contrast: the farm in Vietnam
Cost (US$ kg-1 production) is clearly profitable, while the New Caledonia farm is marginally unprof-
itable under current parameters at a harvest size of 500 g (Fig. 6). This is
b 0.6
because sandfish are unlikely to reach the modelled threshold weight
(~590 g) for profitability in 18 months at New Caledonia using pub-
New Caledonia lished practices (Agudo, 2012; Bell et al., 2007). Both grow-out time
(US$ kg-1 production)

0.4
and component costs were much higher in New Caledonia, with costs
0.2 of purchasing juveniles, labour costs and electricity for pumping being
Net benefit

0 the major drivers of this difference. Farm operating costs and fuel, repairs
and maintenance were more consistent between models. The consistent
-0.2
tidal range on the central coast of Vietnam is an enormous asset to finan-
-0.4 cial viability of pond farming by obviating pumping costs. We modelled
-0.6 larger farms (up to 50 ha) for New Caledonia, but found only minor
economies of scale due to the higher labour costs. A price dividend of
-0.8
470 520 570 620 670 20%, which may be achieved due to consistent, high-quality supply
Weight attained at 18 months (g) would improve prospects for profitability.
The models show that pond farming is a viable alternative to
shrimp for small-scale farmers in Vietnam, corroborated by the actual
Labour expansion of the industry (Duy, 2012; Mills et al., 2012). Conversely,
Juveniles in countries such as New Caledonia with high labour and utility mar-
Operating kets, pond monoculture of sandfish will struggle to be profitable un-
F.O.R.M. less producers can obtain faster growth rates and/or higher market
Electricity prices for the harvested animals than used here (discussed later).
0 0.2 0.4 0.6 0.8 1 1.2 1.4 The price at which sea cucumber juveniles can be produced in the
Cost (US$ kg-1 production) hatchery is also critical for economic viability of farming and sea
ranching (Purcell and Simutoga, 2008). Future improvements in cul-
Fig. 6. Plots of modelled net benefit (price received minus production costs) for 8 ha pond ture technology could allow a reduction in the price of juveniles to
farms in (a) Vietnam and (b) New Caledonia. Net benefit is a function of average growth improve profitability. However, we expect that the economics of pro-
attained over each growing cycle (Vietnam: 10 months; New Caledonia: 18 months). Bar
charts show the component costs for the production of 1 kg of sandfish for sizes readily ac-
ducing sandfish over long culture cycles will direct proponents in
cepted in each country (Vietnam: 350 g; New Caledonia: 500 g). F.O.R.M is fuel, oil, repairs many countries towards co-culture or extensive sea ranching. Exten-
and maintenance. Methodology is given in Appendix S1. sive grow-out modes may also avoid high variation in growth rates
 Author's personal copy
78 S.W. Purcell et al. / Aquaculture 368-369 (2012) 68–81
among animals experienced in intensive pond culture and sea pens
(c.f. Robinson and Pascal, 2012).
5.2. Improving returns
Mariculture is often most profitable when animals are grown only to
the minimum size that markets will accept (Tlusty et al., 2011). Howev-
er, that strategy may not hold if the juveniles are expensive to produce
and if larger animals command higher prices per unit weight — as is the
case for most sea cucumbers. Prices for sea cucumbers locally and on the
Asian dried seafood markets tend to rise exponentially with increasing
body size (Brown et al., 2010; Kinch et al., 2008) (Fig. 7). For example,
based on a conversion of 5% to dried weight from whole fresh weight
(Conand, 1990) and the relationship between retail price and specimen
weight (Fig. 7), a sandfish (H. scabra) of 300 g whole weight is eventu-
ally sold dried (in retail shops) for about US$2.4 whereas a sandfish of
1 kg is eventually sold dried for about US$18. Therefore, we wonder
whether greater profits could be made by allowing tropical sea cucum-
bers to grow to large market size (e.g. 1–1.5 kg) instead of harvesting at
the minimum size that traders will accept.
A further consideration, often overlooked, is that species such as
sandfish have been found to grow at a maximum absolute rate (i.e. g
ind−1 month−1) when they are at an intermediate size of 300–800 g
(Purcell and Kirby, 2006). In extensive sea ranching, it may be possible
to allow the animals 3 to 5 years to reach large sizes that command
‘A+’ grade prices. Ironically, sandfish in Vietnam, Philippines and
Madagascar have usually been harvested at the start, or before, this
peak period of weight gain (Duy, 2012; Juinio-Meñez et al., 2012b;
Robinson and Pascal, 2012). Harvesting sea cucumbers at a small market
size from ponds, sea pens or sea ranches may be dictated by reasons of
cashflow, risks and production turnover. Clearly, a cost-benefit study
across countries is needed to determine the most profitable size for
harvesting tropical sea cucumbers, considering costs, risks and returns,
using established software tools (Johnston, 2012).
One difficulty in comparing yields of sea cucumbers among pub-
lished studies is the disparities in units and metrics. Reports need
700
600
Retail price (US$ kg-1)
500
400
300
200
100
0 in earthen ponds
0 20 40 60 80
Weight (g dried)
4 6 8 10 12 14
Length (cm dried)
Fig. 7. Relationship between retail price of dried sandfish (Holothuria scabra) in shops in
Hong Kong and the weights (blue, squares, solid line) and lengths (red, dots, dashed line)
of individual whole specimens. The best two-parameter models, based on highest r2 values,
were chosen from numerous competing models using DataFit™ 9 software. Price vs weight:
r2 =0.87; equation: y=107.8+(1.29∗x∗ln(x)). Price vs length: r2 =0.82; equation: y=
9647.4∗x(- 13.87/x). Conversion of HK$ prices (listed per 600 g) to US$ was at the international
rate of 0.129. Measurements were made at four shops and 3–4 specimens were sampled ran-
domly from two bins per shop. Total lengths were measured to the nearest 0.5 cm; weights
were measured to ±1 g using an electronic balance. (For interpretation of the references to
colour in this figure legend, the reader is referred to the web version of this article.)
to be clear about the age of the harvested animals, their weights
and whether the animals have been drained of water or gutted
prior to weighing. Profitability comparisons among studies must
also acknowledge the different ‘farm-gate’ or landing prices among
countries (e.g. see Purcell and Simutoga, 2008) due to market struc-
tures and compound supply chains. Farming and ranching of sea cu-
cumbers have the potential to command better prices from buyers for
the raw animals because sizes of the animals can be controlled and
yields per harvest day should be more predictable than wild animals
caught in the fishery. These factors may spur large producers to do the
postharvest processing and exportation themselves and/or develop
markets and supply chains for new product forms (see Perez and
Brown, 2012; Simos, 2012).
6. Conclusions and future opportunities
While important advances have been made in the arenas of larval
culture, nursery systems and grow-out frameworks, few until recently
have been published. The ACIAR-SPC Asia-Pacific Tropical Sea Cucum-
ber Aquaculture Symposium was a landmark forum for identifying
key research priorities and documenting recent advances (Hair et al.,
2012). Sea cucumber mariculture in the tropics is still constrained by
unreliable and generally low survival rates in the hatchery, attributable
in part to few rigorous experimental studies on feeding, stocking densi-
ty and culture systems. Our knowledge base about treatment of disease
and parasites, and the feeding, stocking densities and management of
grow-out in ponds remains poor. A key lesson revealed within and
among studies is that results are not always transferrable from one loca-
tion to another.
Encouragingly, good survival and growth rates have now been
documented in ponds to give promise to profitability where ponds
can be leased or bought cheaply, labour costs are low and seawater
can be exchanged by tidal water movement. The development of
co-culture of sea cucumbers with other taxa struggled in early studies
owing to the choice of partner candidates but still holds great poten-
tial. We anticipate further investment in co-culture, given that sea cu-
cumbers are high-value and a low-trophic-level group that can feed
on the waste products of other mariculture species. Grow-out in sea
pens or in open sea-ranching scenarios will need to contend with
risks of environmental perturbations, predation, poaching and social
conflicts. Sociological issues, such as governance, consultation and
poaching, are significant and must be tackled at the outset.
The key researchable knowledge gaps in the mariculture and
stocking of tropical sea cucumbers discussed here and identified in
the ACIAR-SPC symposium are as follows:
1. Genetic effects of hatchery and stocking practices
2. Broodstock conditioning and egg quality
3. Optimal density of larvae for best settlement per tank
4. Feeding ecology of larvae and optimal microalgae diets
5. Optimal protocols for disease and parasite treatments
6. Stocking density, feeding regimes and water quality management
7. Co-culture opportunities and viability at various sizes of sea
cucumbers
8. Optimal habitats for release into sea pens or for sea ranching
9. Suitable stocking densities in the sea as a function of habitat features
10. Effects of behavioural conditioning and acclimation on post-release
survival of juveniles
11. Strategies for improving governance and communication
12. Comparative analysis of benefits from farming and sea ranching.
Acknowledgements
We thank participants at the ACIAR-SPC Asia-Pacific tropical Sea Cu-
cumber Aquaculture Symposium for inputs and discussions that led to
 Author's personal copy
S.W. Purcell et al. / Aquaculture 368-369 (2012) 68–81
the production of this article. We thank three anonymous reviewers for
their comments on an earlier version of the manuscript. The prepara-
tion of the manuscript was supported by the Australian Centre for Inter-
national Agricultural Research (ACIAR), the WorldFish Centre, Southern
Cross University and James Cook University. For providing information
on commercial hatchery production in various countries, we thank N.
Agudo, J. Altamirano, P. Asha, G. Robinson, I. Eeckhaut, T. Lavitra, B.
Pascal, N. Duy, P. Becker, R. Costa, B. Giraspy, A. Mercier, M. Ito and W.
Bowman. Data on retail prices in Hong Kong were gained from project
2010–004 of ACIAR's PARDI initiative with the support of W. Seng Wei
and R. Ram.
Appendix A. Supplementary data
Supplementary data to this article can be found online at http://
dx.doi.org/10.1016/j.aquaculture.2012.08.053.
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