F I S H and F I S H E R I E S , 2013, 14, 3459

Sea cucumber fisheries: global analysis of stocks,

management measures and drivers of overfishing

Steven W Purcell1, Annie Mercier2, Chantal Conand3, Jean-Francois Hamel4, M Veronica Toral-Granda5,
Alessandro Lovatelli6 & Sven Uthicke7

1
National Marine Science Centre and Marine Ecology Research Centre, Southern Cross University, PO Box 4321, Coffs
Harbour NSW 2450, Australia; 2Ocean Sciences Centre, Memorial University, St. Johns NL, A1C 5S7, Canada;
3
Laboratoire dEcologie Marine, Universite de La Reunion, PO Box 7151, 97715 Saint Denis, Reunion, France and MNHN,
57 rue Cuvier, 75005 Paris, France; 4Society for the Exploration and Valuing of the Environment (SEVE), 21 Phils Hill
Road, Portugal Cove-St. Philips NL, A1M 2B7, Canada; 5World Wildlife Fund (WWF), Galapagos Programme, PO box
20-10-10, Puerto Ayora, Santa Cruz Island, Galapagos, Ecuador; 6Fisheries and Aquaculture Resources Use and
Conservation Division, Food and Agriculture Organisation of the United Nations, Viale delle Terme di Caracalla, 00153
Rome, Italy; 7Australian Institute of Marine Science, PMB No. 3, Townsville Qld 4810, Australia

Abstract Correspondence:
Worldwide, most sea cucumber fisheries are ineffectively managed, leading to Steven W. Purcell,
National Marine
declining stocks and potentially eroding the resilience of fisheries. We analyse trends
Science Centre,
in catches, fishery status, fishing participation and regulatory measures among 77 Southern Cross
sea cucumber fisheries through data from recent fishery reports and fishery University, PO Box
managers. Critical gaps in fisheries biology knowledge of even commonly targeted 4321, Coffs Harbour,
species undermine the expected success of management strategies. Most tropical NSW 2450, Australia
Tel.: +61 2 6648
fisheries are small-scale, older and typified by numerous (>8) species, whereas
3900
temperate fisheries are often emerging, mono-specific and industrialized. Fisher Fax: +61 2 6651
participation data indicated about 3 million sea cucumber fishers worldwide. Fisher 6580
participation rates were significantly related to the average annual yield. PERMANOVA E-mail: steven.w.
analysis showed that over-exploited and depleted fisheries employed different sets of purcell@gmail.com

measures than fisheries with healthier stocks, and a non-metric multidimensional Received 2 May 2011
scaling ordination illustrated that a broad set of regulatory measures typified Accepted 30 Sep 2011
sustainable fisheries. SIMPER and regression tree analyses identified that the
dissimilarity was most related to enforcement capacity, number of species harvested,
fleet (vessel) controls, limited entry controls and rotational closures. The national
Human Development Index was significantly lower in countries with over-exploited
and depleted fisheries. Where possible, managers should limit the number of fishers
and vessel size and establish short lists of permissible commercial species in
multispecies fisheries. Our findings emphasize an imperative to support the
enforcement capacity in low-income countries, in which risk of biodiversity loss is
exceptionally high. Solutions for greater resilience of sea cucumber stocks must be
embedded within those for poverty reduction and alternative livelihood options.

Keywords Beche-de-mer, ecosystem approach, fisheries stocks, invertebrate,

regulatory measures, resource management

34 DOI: 10.1111/j.1467-2979.2011.00443.x 2011 Blackwell Publishing Ltd

 Management of sea cucumber fisheries S W Purcell et al.

Introduction 35
Biological challenges for management 37
Insufficiency of life-history parameters 37
Ecological vulnerability 37
Ecosystem effects of fishing 38
Data collection and methods 38
Status of sea cucumber fisheries 41
Small-scale fisheries 42
Fishing methods 43
Processing methods and trade 43
Cultural and socio-economic drivers 43
Subsistence uses and subsistence fishing 44
Industrialized, highly capitalized fisheries 44
Emerging fisheries 44
Fishing methods 44
Processing methods and trade 45
Trends in sustainability and exploitation 45
Fisher participation 45
Management measures 46
Human development and sustainability 48
Discussion 49
Enforcement drives sustainability 49
Management measures 50
Poverty, participation rates and fisheries sustainability 50
Measures to mitigate loss of biodiversity 51
Pulse vs. continuous fishing 52
Conclusions 53
Acknowledgements 54
References 54
Supporting Information 59

genic stressors and victims of under-funding and

Introduction
With more than half of the worlds finfish fisheries
at their maximum sustainable limit and 28 per cent
over-exploited, depleted or recovering (FAO 2010a),
increased pressure is being shifted to non-finfish
resources (Pauly et al. 1998; Anderson et al.
2011a). Whether a declining mean trophic level of
fisheries landings is attributed to serial replacement
of high-trophic-level fisheries with low-trophic-level
fisheries (Pauly et al. 1998, 2002) or simply the
serial addition of low-trophic-level fisheries (Essing-
ton et al. 2006), exploitation of low-trophic-level
invertebrate resources is occurring at a faster rate
than management can evolve (Anderson et al.
2011a, b). Many of the newly targeted invertebrate
resources occur on tropical coral reefs, which are
vulnerable to various direct and indirect anthropo-
under-management (Sadovy 2005). Less certain are
the potential flow-on effects of overfishing inverte-
brates to ecosystem functioning (Anderson et al.
2011b) and whether the adoption of an ecosystem
approach to management (sensu Garcia et al. 2003)
can rebuild stocks and stabilize boom-and-bust
oscillations in these fisheries.
Throughout the world, 66 species of sea cucum-
bers are commonly exploited (Purcell 2010) and of
particular importance in the Indo-Pacific (Conand
and Byrne 1993; Choo 2008a; Conand 2008;
Kinch et al. 2008a). In Australia, for example, they
appear to have been the first export commodity,
with involvement of Aboriginals pre-dating Euro-
pean explorers (Macknight 1976; Russell 2004;
Manez and Ferse 2010). Sea cucumbers have been
harvested for Chinese consumers for at least

2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459 35

Management of sea cucumber fisheries S W Purcell et al.

400 years (Akamine 2001; Manez and Ferse 2010) 2007). Resilience of socialecological system (SESs)
and have been economically important in the is an imprecise concept (Walker et al. 2004) aligned
western Pacific for over a century (Conand 1990; with adaptive capacity (Carpenter et al. 2001).
Kinch et al. 2008a). In New Caledonia and Tonga, Here, it concerns the ability of fishing communities
sea cucumber export earnings recently surpassed to detect hard-to-reverse thresholds (Lebel et al.
finfish and other marine resources (Purcell et al. 2006) and adapt management to changes in stocks
2009a; P. Ngaluafe, Ministry of Fisheries, Tonga, such that the functionality of the SES is maintained
pers. comm.) and elsewhere are the leading inver- (Walker et al. 2002, 2004; Andrew et al. 2007).
tebrate fishery export (Anderson et al. 2011b). Resilience in sea cucumber fisheries requires partic-
During the 1980s and 1990s, more countries ipatory management systems that combine local
started exporting sea cucumbers (Conand 1997, and scientific knowledge (Berkes and Seixas 2005)
1998) and global production boomed (Fig. 1). Sea and enable fishers, processors and exporters to take
cucumber fisheries are active, or recently so, in at responsibility for the resource (Andrew et al. 2007).
least 70 countries from all of the worlds oceans Exploitation must then be adapted such that stocks
(reviews in Toral-Granda et al. 2008). are not depleted to states that compromise their role
Small-scale fisheries, like many centred on coastal to ecosystems and livelihoods. However, engaging
invertebrates, are important in more than 90% of countless small-scale fishers in low-income coun-
the worlds EEZs and are increasingly predominant tries is arduous (Conand and Muthiga 2007; Kinch
in low-income countries (Mora et al. 2009). They et al. 2008b; Ochiewo et al. 2010).
are harder to assess and poorly reported because Most sea cucumber fisheries have a boom-and-
they involve many times more fishers than indus- bust history (Conand 1990; Uthicke 2004; Uthicke
trialized fisheries (Berkes et al. 2001; Worm et al. et al. 2009). The decimation of stocks has forced
2009). As a mainstay or livelihood supplement, sea recent moratoria in Commonwealth of the Northern
cucumber fisheries are significant to poverty reduc- Mariana Islands, Costa Rica, mainland Ecuador,
tion in developing countries (Conand and Muthiga Egypt, India, Mauritius, Mayotte (France), Panama,
2007; Conand 2008; Kinch et al. 2008a, b). Papua New Guinea, Solomon Islands, mainland
An ecosystem approach to fisheries (EAF) man- Tanzania, Tonga, Vanuatu and Venezuela (Purcell
agement with a focus on stakeholder participation 2010). The moratoria have sometimes given rise to
and the need to foster resilient fisheries have been illegal harvesting (Conand 2006). Many sea cucum-
strongly advocated (Garcia et al. 2003; Hilborn ber fisheries are precarious (sensu Walker et al.
2004; Berkes and Seixas 2005; Andrew et al. 2004) with respect to resilience of the SESs.
A number of species are threatened and there is
Global sea cucumber production (t dried)

evidence of local extinctions in some tropical regions

20000 Aquaculture production
Capture fisheries (Uthicke and Benzie 2001; Uthicke and Conand
2005; Friedman et al. 2011). Urgently needed are
15000 radical different management paradigms and new
instruments to safeguard the reproductive capacity
10000 of stocks.
Recent workshops have striven to build a plat-
5000 form of current information on which new man-
agement systems can develop (Lovatelli et al. 2004;
0
Bruckner 2006; Friedman et al. 2008). Likewise,
1950 1960 1970 1980 1990 2000 new reviews now depict the status of sea cucumber
Year fisheries in regions around the world (Choo 2008a;
Conand 2008; Hamel and Mercier 2008a; Kinch
Figure 1 Global wild captures and aquaculture produc-
et al. 2008a; Toral-Granda 2008a). Anderson et al.
tion of sea cucumbers over time; in metric tonnes (t).
(2011b) showed the fast boom-and-bust rate of 23
Source FAO Fishstat, with correction: data for aquaculture
production and wild captures of Apostichopus japonicus
large sea cucumber fisheries and likely drivers. Still
(China, Japan, Korea) refer to fresh animal weights, so lacking is a global appraisal of stocks across all
these were converted to dried weight using a conversion fisheries, examination of fishing participation and
factor of 0.04, based on other Stichopus species (Skewes yields to fishers, and a critical examination of
et al. 2004). potential management solutions.

36 2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459


Here, we present the first meta-analysis of the
state of nearly all global sea cucumber fisheries and
their regulatory measures. We highlight three key
biological challenges facing the management of sea
cucumber fisheries through a review of recent
research. New data were compiled from published
sources and agencies from 77 fisheries from the
main fishing regions of the world: Africa and the
Indian Ocean, South-East Asia, the Western Cen-
tral Pacific, the Northern Atlantic and Pacific, and
Latin America and the Caribbean. We use both the
data and reviews of recent published reports to
describe the state of fisheries. We further examine
participation rates of fishers and analyse the
relationship of variables to stock status across the
global data set. Our findings lead us to propose a
radical shift in management paradigms within an
EAF that could reverse the gloomy prognoses for
certain sea cucumber fisheries.
Biological challenges for management
Insufficiency of life-history parameters
A fundamental barrier to improved management of
sea cucumber fisheries is the lack of data on basic
biological parameters of most species (Conand
1989; Conand and Sloan 1989; Lovatelli et al.
2004). Their soft bodies preclude most tagging
methods for long-term markrecapture studies to
assess growth, movement, mortality rates and
longevity. Genetic fingerprinting (Uthicke and
Benzie 2002) and fluorochrome marking of ossicles
(Purcell et al. 2006; Purcell and Blockmans 2009)
are the only reliable long-term markers to date and
both have limitations. Larvae cannot yet be easily
identified to species level for quantifying dispersal
and settlement events, and the cryptic nature of
juveniles of the many reef-associated species obvi-
ates direct estimates of recruitment. In addition,
analysis of cohorts is confounded because juveniles
from the same cohort can grow at hugely variable
rates (Hamel and Mercier 1996; Purcell and Sim-
utoga 2008; So et al. 2010) and adults can shrink
in size (Hamel and Mercier 1996; Uthicke et al.
2004). Classical fisheries science approaches based
upon such parameters are therefore tenuous for
most commercial sea cucumber species (Conand
1990; Purcell 2010).
Similarly, more contemporary management
strategies, like rotational harvest strategy, also
struggle with the lack of biological data on most
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459
Management of sea cucumber fisheries S W Purcell et al.
exploited sea cucumbers. In Alaska (USA), British
Columbia (Canada) and Queensland (Australia),
rotational harvest closures have been developed to
spread fishing effort spatially (Hamel and Mercier
2008a; Purcell 2010). The 3-year harvest cycles are
linked to species quotas but inherently assume that
growth rates and recruitment rates of the fished sea
cucumbers are equally fast. These premises are
doubtful for at least some key tropical (Uthicke et al.
2004) and temperate species (Hamel and Mercier
1996; So et al. 2010).
Ecological vulnerability
Although the deeper habitats of certain temperate
species constrain fishing effort in dive and trawl
fisheries (Hamel and Mercier 2008a), most tropical
species tend to occur in shallow water within the
range of breath-hold diving (Kinch et al. 2008a).
Displacement rates are also sluggish (Conand 1991;
Purcell and Kirby 2006; Shiell and Knott 2008), so
sea cucumbers are incapable of moving away
quickly from patches of high density once detected
and exploited by fishers.
As gonochoric broadcast spawners, sea cucum-
bers need to be in close proximity to mates for
successful fertilization of gametes (Babcock et al.
1992; Shepherd et al. 2004; Mercier and Hamel
2009). In low-density populations (e.g. a few
individuals per ha), they may fail to get close to
conspecifics in breeding periods, resulting in asyn-
chronous spawning (see review by Mercier and
Hamel 2009). Research is still needed to determine
minimum population densities for positive rates of
per capita population growth (Lovatelli et al. 2004;
Bell et al. 2008; Friedman et al. 2011). In addition,
high-value species on tropical reefs often co-habit
with those of low-value, allowing opportunistic
depletion of high-value species by fishers after
shifting to low-value species.
High longevity in commercial species is related to
slow recovery rates and makes stocks more vulner-
able to collapse (Caddy and Seijo 2005). Perhaps a
few species are short (but still several years to a
decade) lived (e.g. S. chloronotus; Conand 1989),
with regular recruitment. On the other hand,
Conand (1989) modelled that A. mauritiana, A. echi-
nites and Thelenota ananas lived at least 1015 years,
and these values are now recognized as underesti-
mates because we now know that adult sea
cucumbers can live several decades and shrink in
size (Uthicke et al. 2004). Similarly, the cold-
37
Management of sea cucumber fisheries S W Purcell et al.
temperate Cucumaria frondosa appears to require
1025 years to attain full adult size and is probably
quite long lived (Hamel and Mercier 1996; So et al.
2010). Low recruitment (Uthicke 2004; Hearn et al.
2005), slow growth and high longevity may render
some species as k-strategists and thus especially
vulnerable to overfishing (Conand 2001).
Ecosystem effects of fishing
Overfishing of sea cucumbers could also lead to
indirect, deleterious effects on reef and soft-bottom
ecosystems. The juveniles are important prey in food
webs (Francour 1997; So et al. 2010), and adults are
important in ecosystem function (Birkeland 1988;
Uthicke 2001b; Wolkenhauer et al. 2010). For
example, some species regularly bury, thereby oxy-
genating sediments through bioturbation (Bakus
1973; Massin 1982; Uthicke 1999, 2001a; Purcell
2004). Many aspidochirote sea cucumbers consume
large quantities of sediments (Uthicke 1999) and
convert organic detritus into animal tissue and
nitrogenous wastes, which can be taken up by algae
(Uthicke and Klumpp 1998; Uthicke 2001b) and
seagrasses (Wolkenhauer et al. 2010) and increase
their productivity, thus producing more available
food for herbivores. On coral reefs, where nutrients
may be limiting, this role in nutrient recycling is
likely to be significant (Uthicke 2001b). In addition,
removal of sea cucumbers in aquarium experiments
has resulted in developments of algal mats, which
may have negative effects on sediment infauna as a
result of potential anoxia (Uthicke 1999). Removal
of sea cucumbers in some systems may therefore
reduce primary production for the whole food chain
and affect sediment infauna by reducing the aerobic
layer of sediments. In temperate waters, dendrochi-
rote sea cucumbers (e.g. Cucumaria frondosa) may be
especially important in recycling suspended matter
(Hamel and Mercier 1998). Management plans we
reviewed rarely addressed the risks to ecosystems or
weighted regulatory measures on the ecological
services of certain species to ecosystem function. For
instance, managers could list and rank vulnerability
of other locally exploited soft-bottom or herbivorous
species to indirect impacts from the removal of sea
cucumbers. Additionally, sea cucumber species that
bioturbate sediments vigorously (e.g. Holothuria
scabra, H. lessoni, Actinopyga spinea, Bohadschia viti-
ensis) could have relatively larger minimum size
limits or be excluded from the list of species that can
be exploited.
38
Data collection and methods
Fisheries are management units that include the
stock and fishers and, for the purpose of the data set,
we considered multispecies fisheries as a fishery. We
were unable to obtain reliable data for a small
number of countries (<10) in which we were aware
that sea cucumbers were being, or have recently
been, fished so we excluded those countries (e.g.
Italy, Greece, North Korea, Somalia, Spain, Yemen).
Data from 77 sea cucumber fisheries from 62
countries were collated from recent regional reviews
(Toral-Granda et al. 2008), national fisheries assess-
ment reports, recently published accounts and
direct correspondence with senior fisheries officers
and national fisheries specialists (Table S1). Fisher-
ies were classed as follows: small-scale if the
predominant mode of fishing involved traditional
gear with small vessels (following Berkes et al.
2001); industrialized if large vessels and modern
fishing gears were mainly used; or a combination of
the two if the scales of fishing presented in similar
frequency in the fishery. We categorized illegal
fisheries as those in which fishing was prohibited
(i.e. by moratorium) but sea cucumbers were still
being collected and sold by fishers on the black
market; these did not include fisheries in which
some illegal fishing activities or unreported fishing
occurs among legal fishing operations. Some sub-
sistence fishing of sea cucumbers occurs in many
fisheries, so we defined subsistence as those fisheries
in which harvesting was solely for domestic con-
sumption (i.e. no exportation).
Fishery production in dry weight (beche-de-mer
or trepang) were averaged over the period 2004
2008 for each fishery, based on FAO data of exports
(FAO Fishstat), regional reports and data solicited
from fisheries agencies. Because of a lag in trade
reporting, this was the most recent data available at
the time of analyses. Sea cucumbers in most
countries are practically solely exported dried
(Ferdhouse 2004; Kinch et al. 2008a), whereas
Apostichopus japonicus from South Korea, Japan and
China is often sold undried, and FAO catch (i.e.
export) data for these fisheries mostly reflects wet
weight (Conand 2001; Akamine 2004). Therefore,
we used 0.04 as an approximate conversion for
catch data from those countries (based on other
Stichopodid species; c.f. Skewes et al. 2004; Purcell
et al. 2009b). In a few cases, we used data on Hong
Kong imports because these could not be obtained
from national export records. We used data from
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459

just one or more years if these were the only
available in this 5-year period. Some fisheries
produced exports for some years and then were
closed by moratorium, so we analysed the produc-
tion data and regulatory measures from the years
of production but also denote these as having
moratoria. Gutted fresh weights or whole animal
weights, for example for production statistics in
fisheries reports from Coral Sea and Torres Strait
(Australia), were converted to dry weights using a
conversion ratio of 0.158 for mixed reef species
harvested (Skewes et al. 2004; Purcell et al. 2009b).
As in other fisheries, taxa with highest potential
profit are targeted first (Sethi et al. 2010). We
ascribed the exploitation status of fisheries based on
(i) the extent of change in annual exports of high-
and medium-valuable species (following Annex
10.1 in Purcell 2010) over the past two decades,
or overall decline in exports in mono-specific fish-
eries or where data for species were grouped, and/or
(ii) the extent of change in abundances of high- and
medium-value species in the wild from fishery-
independent surveys. We used fishery-independent
data (e.g. from underwater visual census) where
available. In other cases, we used export data,
which will have an inherent underestimation
of exploitation state in many instances because
these statistics are not often separated by species
(Ferdhouse 2004; Choo 2008b). This is problematic
because lower-value species will be targeted by
fishers, at the same or higher rate, when high-value
species become scarce (Kinch et al. 2008a; Ander-
son et al. 2011b). That is, export volume may
increase from a country as stocks of valuable species
are diminished because fishers resort to collecting
greater volumes of low-value species that may be
relatively more abundant. Where possible, abun-
dances in fishing grounds were compared to virgin
stock abundance in permanent no-take zones in
comparable habitats. We recognized that assigning
one status category for multiple species in a fishery
was problematic but this was necessary because the
stock status for individual species in fisheries was
usually not possible and we needed a measure by
which to compare fisheries and relate management
measures. Our exploitation status categories follow
standard definitions by FAO (2005, p. 213) and
adaptation of criteria by Sumaila et al. (2007):
Under-exploited: Undeveloped; believed to have a
significant potential for expansion in total produc-
tion. Average recent annual catches (usually based
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459
Management of sea cucumber fisheries S W Purcell et al.
on exports) at least 50% of the peak maximum
potential catch, which is not yet realized. Popula-
tion abundance is at least 90% of virgin abundance.
Moderately exploited: Exploitation rate is sustain-
able for medium- and high-value species, with
some potential for increased yields without
diminishing the reproductive capacity of stocks.
Annual catches are at least 50% of the peak
maximum potential catch, which is not yet
realized. Population abundance is between 60%
and 90% of virgin abundance.
Fully exploited: The fishery is operating at or close
to a maximum sustainable yield, with no
expected room for further expansion. Catches
are increasing or stable and not expected to be
able to increase sustainably. Population abun-
dance is between 30% and 60% of virgin
abundance.
Over-exploited: The fishery is, or has been, exploited
at above a level that is believed to be sustainable
in the long term, with no potential room for
further expansion and a higher risk of stock
depletion/collapse. Annual catch of high- and
medium-value species is between 10% and 50%
of the historical maximum catch, or their abun-
dance is between 10% and 30% of virgin
abundance.
Depleted (collapsed): Catches, and/or stocks, are well
below historical levels, irrespective of the amount
of fishing effort exerted. Catch of medium and
high-value species is below 10% of historical
maximum catch, or their abundance is <10% of
virgin abundance (sensu Worm et al. 2009).
For example, Madagascar ranked as depleted
because annual exports have declined by >85%
from the historical peak but catch is now mostly
small individuals of low commercial value, so
declines of medium- and high-value species are
probably >90% from virgin abundance. The fishery
on the Great Barrier Reef, Australia, ranked as fully
exploited because at least one high-value species
(e.g. Holothuria whitmaei) has been depleted to about
20% of virgin abundance in fishing grounds
(Uthicke 2004), catches of some medium-value
species (e.g. Thelenota ananas) declined by >20%
from previous years (Queensland DPIF 2007), but
stocks of some other medium-value species are
>50% of virgin biomass.
The management system and regulatory mea-
sures used in the period 20042008 within each
fishery, wherever possible, were obtained from
39
Management of sea cucumber fisheries S W Purcell et al.
recent reports, reviews and direct communications
with fisheries agencies. Data on use of regulatory
measures were binomial; 1 was scored if the
measure has been employed, or 0 if not, in the
fishery during the past decade. Capacity controls
were distinguished as fleet controls (on number or
size of boats) and limited entry controls (on number
of fishers). Moratoria were included if harvesting
some or all species was prohibited in the fishery for
at least 1 year (sensu Purcell 2010). Enforcement
capacity was ranked based on number of compli-
ance officers in each fishery compared to the fishery
size and the system for penalizing infringements:
1 = weak, 2 = moderate and 3 = strong. For exam-
ple, Seychelles has strong enforcement capacity
because the managing authority has been able to
license a small number of fishers, their logbooks are
required to be submitted regularly and non-compli-
ance can result in non-renewal of their fishing
licence, sales receipt books are required for sales
between fishers and the processor/exporters, and
random inspections are carried out at processing
facilities and prior to export (Aumeeruddy and
Conand 2008). In contrast, the Republic of Kiribati
has weak enforcement because there is only one
compliance officer who does not physically inspect
exports, fishers catches are never inspected and
rampant exploitation of juvenile sea cucumbers
goes unchecked (K. Tamuera, Ministry of Fisheries,
pers. comm.).
We tested for significant differences (a = 0.05) in
the multivariate suites of measures controllable by
Table 1 Variables potentially controllable by the fishery
manager used in the multivariate analyses. Regulatory
measures are denoted with asterisks.
Variable Data range
Number of species exploited 147
Enforcement capacity 13
Size limits* 01
Seasonal closures* 01
Gear restrictions* 01
Quotas (TACs or ITQs)* 01
Licensing and market chain reporting* 01
Limited entry controls* 01
Fleet controls (boat numbers and sizes)* 01
Permanent no-take reserves* 01
Rotational harvest closures* 01
Moratoria* 01
Number of regulatory measures used 010
40
fishery managers (Table 1) among fisheries with
different status categories using permutational mul-
tivariate analysis of variance (PERMANOVA) in PRI-
MER 6 software with 1000 permutations and using
simple contrasts (Clarke and Gorley 2006). Com-
plete data on regulatory measures (including mor-
atoria), the total number of regulatory measures
used, enforcement capacity and number of species
harvested in each fishery were obtained for 67
(87%) of the sea cucumber fisheries. These variables
could each be potentially controlled by the fishery
manager. We normalized data by z-transformation
(mean = 0, SD = 1) prior to analysis and used
Euclidean distance as the dissimilarity measure.
A SIMPER analysis subsequently examined control-
lable measures contributing to the separation
between groups identified in the PERMANOVA
contrasts (Clarke and Warwick 2001). We chose
non-metric multidimensional scaling (nMDS) for
ordination and applied Pearsons correlation of the
normalized data for vectors of the management
measures. We used the same parameters for the
SIMPER analysis in a complementary regression tree
(RT) analysis in R (R Development Core Team
2010). Regression trees are ideal to investigate
complex ecological data and can handle a variety of
data types including numerical, ranking and rating
data (Death and Fabricius 2000). In the RT
analysis, fishery status was taken as the response
variable and was transformed to categorical data
from 1 to 5 (underexploited to depleted). The optimal
length of the RT was determined by cross-validation
as suggested by Death and Fabricius (2000).
Human Development Index (HDI) rankings for
countries were obtained from UNDP report for 2009.
The calculation for this index incorporates various
measures including poverty, health and education.
We compared average HDI scores across status
categories using a single-factor ANOVA and con-
firmed homoscedasticity of data with Levenes test
using SPSS 17.0 software. Data were non-normally
distributed within groups, and potential transforma-
tions could not improve that, but ANOVA is quite
robust to non-normality especially for data sets like
ours with many treatments (status ranks here) and
samples (Underwood 1997). A StudentNewman
Keuls test was used for post hoc comparisons.
Total number of fishers per year during 2004
2008 was obtained from 53% of fisheries from
sociological survey reports, national statistics sur-
veys and unpublished census data from national
fishing agencies (see Table S1c). For fisheries with
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459

very high participation, data were based on conser-
vative extrapolations from the number of boats in
each fishery or number of sea cucumber fishers per
town or island determined from fishery-dependent
surveys. For example, participation in Philippines
was based on a census total of 469 147 boats
(L. Labe, Philippines Bureau of Fisheries and
Aquatic Resources, unpubl. data) and, conserva-
tively, two fishers per boat. We did not factor in
children who collect sea cucumbers intermittently
(e.g. on school holidays) nor fishers who seldom
collect sea cucumbers. Data of >30 000 fishers in a
fishery were rounded down to the nearest 10 000
values.
As most tropical commercial species inhabit coral
reef environs (Conand 1989), we used total area of
coral reef in the fishery, established from GIS
mapping (Spalding et al. 2001; and regional reports,
for example for Australian fishery areas), as a proxy
for the area of fishing ground in each fishery.
Although some exploited tropical sea cucumbers
inhabit non-coral reef habitats, this proxy should be
relatively consistent among fisheries. We then
calculated participation rates in fisheries, as a
measure of fishers per fishable area, by dividing the
number of fishers in tropical fisheries over the coral
reef area in each fishery. Average annual yield of
dried sea cucumbers per fisher was the average
annual production of the fishery divided by the
number of fishers, and we recognize that in many
fisheries some fishers catch disproportionately more
sea cucumbers than other fishers. Initial data
exploration to describe the relationship between
average annual yield and participation rate showed
40E 60E 80E 100E 120E 140E 160E 180 160W 140W 120W
70N
50N
30N
10N
10S
30S
50S
70S
40E 60E 80E 100E 120E 140E 160E 180 160W 140W 120W
Figure 2 Current status of sea cucumber fisheries in global fisheries (n = 69).
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459
Management of sea cucumber fisheries S W Purcell et al.
evidence of a mean/variance relationship. Log2
transformations of both x- and y-axes data
improved this relationship in linear regression and
gave a good fit to the data. We modelled the
relationship between average annual yield per fisher
and participation rate using R software (R Develop-
ment Core Team 2010).
We examined the likely pros and cons of pulse
fishing vs. continuous modest fishing strategies for
sea cucumbers through case-studies of pulse fishing.
Our comparison of likely responses of sea cucumber
stocks to either strategy also considers recent
biological studies on longevity, larval supply and
population turnover rates. A closure-pulse fishing-
closure strategy (sensu Friedman et al. 2011) allows
fishing for one or several years across the fishery
followed by an inactive period of several or many
years. In contrast, seasonal closures are for part of a
year and rotational closures permit fishing within
part of the fishing grounds for short (e.g. 1 year)
periods followed by a specified fallow period (e.g. 2
or 3 years)(Purcell 2010).
Status of sea cucumber fisheries
Our global analysis revealed an alarmingly high
incidence of over-exploitation and depletion of sea
cucumber stocks, particularly in the Indo-Pacific
(Fig. 2). Overall, 20 per cent of fisheries were
depleted and 38 per cent were over-exploited. 14
per cent of the fisheries were fully exploited, with no
potential room for further expansion. Only 27 per
cent of global fisheries were underexploited or
moderately exploited and many of these were in
100W 80W 60W 40W 20W 0
70N
50N
30N
10N
10S
30S
Depleted
Over exploited
Fully exploited 50S
Moderately exploited
Under Exploited 70S
100W 80W 60W 40W 20W 0
41
Management of sea cucumber fisheries S W Purcell et al.
deeper, temperate waters or the fisheries were in
moratoria hiatus or developmental phases.
Small-scale fisheries
The majority (66%) of sea cucumber fisheries
involved small-scale fishing operations for export
(Fig. 3). This seems related to the fact that sea
cucumbers and these fisheries are generally more
common in shallow habitats of tropical waters
where fishing units, time commitment, equipment,
boats and catches at a small-scale (sensu Berkes
et al. 2001) are viable. Small-scale here concerns
the simplicity of fishing gears and boats and
artisanal nature in which animals are collected,
Figure 3 Scale of fishing in global sea cucumber fisheries (n = 77).
Figure 4 Bubble plot of the number of species harvested in different fisheries. The area of each dot corresponds to the
number of species exploited.
42
not the number of fishers, overall catch in the
fishery nor geographic scale of the fishery (sensu
Berkes et al. 2001). Of those fisheries, 27% were
operating illegally; that is, sea cucumbers were still
being harvested and exported despite national
moratoria. The illegal fisheries (official moratorium
but clandestine fishing) occurred in Central and
South America and the Western Indian Ocean
(Fig. 3).
Multispecies fisheries for sea cucumbers are the
norm in the tropics (Conand and Sloan 1989;
Conand and Byrne 1993; Choo 2008a; Conand
2008; Kinch et al. 2008a). Small-scale sea cucum-
ber fisheries, especially those in countries with low
per capita income, concern a large number of
No. species
harvested
=1
= 10
= 20
= 30
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459

species (Fig. 4). Countries within the Coral Triangle,
Melanesia, and parts of the Indian Ocean export
more than 20 different sea cucumber species (Choo
2008a; Kinch et al. 2008a), and there are now 47
species exploited in the Philippines (Labe 2009). In
stark contrast, an average of 4 sea cucumber species
is exploited in Central and South American fisheries
and some are yet undescribed taxonomically.
Although some have well-defined habitat specifici-
ties, as many as a dozen or more tropical species can
occupy the same general habitat and so are
collected together by fishers opportunistically.
Only six of the 55 small-scale sea cucumber
fisheries target just one or two species. The fishing,
handling and processing methods are similar for
various species, with few exceptions. In the Gala-
pagos Islands and mainland Ecuador, Isostichopus
fuscus is the sole species collected legally but fishing
is at a semi-industrial scale because relatively large
boats are used (Toral-Granda 2008b). Only five of
the 18 central-western Pacific island fisheries have
exported fewer than 10 species, and those cases
appear attributable to low productivity of isolated
reef systems and relatively low diversity of stocks
rather than preference by fishers (Kinch et al.
2008a). This trend contrasts starkly with mono-
specific fisheries in many temperate fisheries
(discussed later).
Fishing methods
A unifying trait of most small-scale fisheries is the
simplicity at which sea cucumbers are collected. In
some cases, they are harvested mainly by gleaning
on reef flats and sand flats at low tide. Women and
children play significant roles in sea cucumber
fisheries through gleaning at nearshore sites, for
instance at Cook Islands, Indonesia, Madagascar,
New Caledonia, Oman, the Philippines and Zanzibar
(Choo 2008a,b, Conand 2008; Kinch et al. 2008a;
Purcell et al. 2009a; Eriksson et al. 2010). More
often, sea cucumbers are collected by hand in
fisheries by a combination of gleaners and divers,
using either simple skin-diving equipment (Conand
2008; Purcell et al. 2009a) or surface supplied air
(hookah) and SCUBA from small motor boats or
traditional canoes (Akamine 2001; Eriksson et al.
2010; Ochiewo et al. 2010). The minimal buy-in
costs to harvest sea cucumbers means that it is easy
for smallholders to participate in fishing and there
may be a high number of occasional fishers because
there is no capital rent (Ochiewo et al. 2010).
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459
Management of sea cucumber fisheries S W Purcell et al.
Processing methods and trade
Post-harvest processing in small-scale fisheries is
carried out with simple equipment; practically all
that is needed is a knife, fuel, a large pot and a
drying rack (Brown et al. 2010). The sea cucumbers
are gutted through a small slit in the animals, then
boiled and dried in the sun or over a fire (Conand
1990; Brown et al. 2010). The use of mangrove and
forest wood for fires in processing is a particularly
serious problem requiring management interven-
tion (Purcell 2010). In localities close to urban
centres, sea cucumbers collected artisanally are
more and more frequently sold to middlemen or
exporting companies that process the animals using
modern equipment (e.g. gas stoves and drying
ovens) (Conand 2008; Purcell et al. 2009a). Cen-
tralized processing of this sort probably leads to
more consistent and higher-quality product but to
the loss of value-adding within villages (Friedman
et al. 2008).
Trade chains in small-scale fisheries tend to be
longer and more complex than those of industrial-
ized fisheries. Fishers may sell to village buyers, who
in turn sell the product to middlemen who sell to
exporting agents (Brown et al. 2010; Eriksson et al.
2010). This sea cucumber fishery system is com-
mon throughout the Indo-Pacific (Conand 1997,
1998). Policing trade chains and sales of beche-
de-mer (the dried product, also called trepang) is
therefore difficult in developing nations (Ochiewo
et al. 2010).
Cultural and socio-economic drivers
A further trait of small-scale fisheries are the long-
serving cultural associations with sea cucumber
resources. In the Indo-Pacific, sea cucumbers have
been collected for more than 150 years and have a
special place in social cultures (Conand 1990, 2008;
Akamine 2001; Kinch et al. 2008a). Involvement of
Australian Aboriginals in sea cucumber fisheries
dates to at least 1700s and has been illustrated in
cave paintings, centuries old (Macknight 1976;
Russell 2004). Fishers in the Coral Triangle and
East Africa have been found to rely on sea cucum-
bers for a large, or principal, proportion of household
income (Choo 2008a,b; Kinch et al. 2008b; Ochiewo
et al. 2010). Unlike recent small-scale tropical fish-
eries, like those for marine ornamentals, sea urchins
and live reef fish, management regulations to limit
the rights to fishing sea cucumbers can be an affront
to tradition and generational livelihood activities.
43
Management of sea cucumber fisheries S W Purcell et al.
Subsistence uses and subsistence fishing
Just 4 of the 77 fisheries were solely for subsistence
and local consumption [Cook Islands, Samoa
(Western), Guam, Nauru] (Fig. 3). In other locali-
ties, there is some subsistence fishing but the vast
majority of harvested sea cucumbers are exported,
so we classed those as export fisheries. Certain
species are eaten within some communities in at least
ten western Pacific island nations, and the viscera
(gonads and intestines) are eaten as a delicacy by
some Polynesians (Kinch et al. 2008a). In Fiji, Tonga
and Papua New Guinea, the body wall of the animals
is sold locally and eaten particularly in times of
economic hardship (Kinch et al. 2008a,b). Some
Malaysian communities eat certain sea cucumbers
or use them in medicinal tonics (Choo 2008a), and
they are consumed in some places in the Philippines
(Choo 2008b) and Peru (Toral-Granda 2008a).
Subsistence use is not restricted to the tropics.
Indigenous peoples of western North America and
Arctic Canada have collected and eaten them for
centuries, and they are common in the diets of a
majority of households in some regions (Hamel and
Mercier 2008a). Overall, subsistence fisheries
appear to have lower exploitation rates per fisher
than commercial fisheries.
Industrialized, highly capitalized fisheries
Industrialized fishing, involving large vessels and/
or sophisticated gears, occurred in 25% of fisheries
we assessed. From data at hand, industrialized and
small-scale fishing methods were used at compara-
ble frequency in just three fisheries (Eritrea, Mad-
agascar and New Caledonia) (Fig. 3). In other
cases, such as the Philippines, where some larger
boats are used in collecting sea cucumbers (Ak-
amine 2001), these appear to contribute to a small
proportion of the overall catch and collection gears
are artisanal. In high-income countries, especially
in temperate regions, fisheries tended to be indus-
trialized (Fig. 3). These fisheries differ in multiple
aspects from their small-scale counterparts (Toral-
Granda et al. 2008).
There are generally far fewer fishers in industri-
alized sea cucumber fisheries, making participatory
approaches to management more practical and
presumably facilitating enforcement and the moni-
toring of catches. For example, there are only two
companies involving a handful of boats in the fishery
on Australias Great Barrier Reef, which stretches
more than 2 000 km of coastline. Our data showed
44
that just 15 fishers operate the trawl fishery for
Cucumaria frondosa in eastern Canada and just a few
fishers trawl that species in Iceland. Similarly, <200
divers collect the red sea cucumber Parastichopus
californicus in Alaska (Hamel and Mercier 2008a).
Most of the industrialized sea cucumber fisheries
are relatively young (Anderson et al. 2011b). Sea
cucumbers in many high-income countries are
harvested to diversify commodities as finfish stocks
have been overfished (Anderson et al. 2011a). For
example, commercial landings of sea cucumbers
were first recorded in the early 1980s on the east
coast of Canada and from 1971 along the coast of
Maine, USA (Hamel and Mercier 2008a). Conse-
quently, management agencies do not have to
grapple with generational traditions of fishing these
resources, as occurs in many small-scale fisheries in
the tropics. Nevertheless, some indigenous peoples
in Canada have ancestral rights to subsistence
fishing (Hamel and Mercier 2008a).
Emerging fisheries
Further, some sea cucumber fisheries are still in the
early stages of development. These emerging fisher-
ies epitomize sources of the phenomenon of declin-
ing mean trophic level of global fisheries (Pauly
et al. 1998). However, recent analyses indicate that
mean trophic level of fisheries should not be inferred
from fishery data alone (Branch et al. 2010) and
that many fisheries are also ruled by ease of access
or monetary value (Sethi et al. 2010). Emerging sea
cucumber fisheries have also created new markets
for temperate species and body organs (Hamel and
Mercier 2008a). In other cases, new fisheries have
been spurred by Asian traders looking for new
species and fishing grounds (e.g. Muthiga et al.
2007; Toral-Granda 2008a,b). In Iceland, where
only one licence has been issued, stocks appear to be
limited and the fishery is in a developmental stage
(Hamel and Mercier 2008a). Harvesting of sea
cucumbers in industrialized fisheries has also
emerged in Spain (Balearic Islands) (Ramon et al.
2010), Italy (A. Lovatelli, personal observation),
and other parts of the Mediterranean and Aegean
Sea (see Kazanidis et al. 2010; Antoniadou and
Vafidis 2011) but, owing to a lack of documentation
on status and management, are not described here.
Fishing methods
Most of the industrialized fisheries of North Amer-
ica, Iceland, Russia and Japan are mono-specific
(Fig. 4). The animals are collected either by hand by
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459
 Management of sea cucumber fisheries S W Purcell et al.

divers using surface supplied air (hookah) or using companies support that assertion. Moreover, grad-
scallop or urchin drags or modified trawl gears. ing and packaging of the product prior to export is
Dragnets in Japan comprise a weighted metal frame sometimes achieved by high-tech equipment in
of several metres wide, which is dragged along the industrialized fisheries, which boosts the export
benthos behind large vessels to collect Apostichopus price and reputation. Sea cucumbers are becoming
japonicus (Purcell 2010). In Newfoundland and increasingly exported as frozen or vacuum-packed
Labrador (Canada), Cucumaria frondosa is harvested product by some countries (e.g. Australia, Canada,
from soft-bottom or gravely grounds in an emerging Iceland). Russia now produces hydrated canned sea
fishery using modified drag gears (Hamel and cucumbers (Hamel and Mercier 2008a), and the
Mercier 2008b). For those fisheries using mobile muscle bands and aquapharyngial bulbs are
fishing gears, the issue of habitat destruction is removed from sea cucumbers in Canada and sold
especially important in an EAF. separately from the body wall (Hamel and Mercier
2008a).
Processing methods and trade
Processing steps of most industrialized fisheries are
Trends in sustainability and exploitation
comparable to those in small-scale fisheries; the
animals are gutted, boiled and then dried into
Fisher participation
beche-de-mer (or trepang). However, the equip-
ment used to process the animals is usually more Our data revealed that just over 2 million fishers
sophisticated, such as automated boilers and drying have been recently engaged in collecting sea
ovens. Fishing companies contend that their pro- cucumbers from 39 of the 77 fisheries we examined.
cessing techniques and equipment produce a more In view that many of the 38 fisheries for which the
consistent high-quality product than small-scale number of fishers were not obtained were in Asian
fishers, and the high export prices of some of those and Latin American countries (including mainland

1000 Moderately exploited

Av. yield per fisher (kg dried per year)

Ag Nc
Fully exploited
800 An Over exploited
Depleted
Fp Ac
600
Er
400
Cu Se Mu
Aw
200 Ta At To
Wa My Mg
Gu Ke Kb Pn Sa
Eg Om In
0 Va Sr Gp Ph
Ma

1 0 1 2 3 4 5 6 7 8
Participation rate (log2 (No. fishers 100-km2)

Figure 5 Relationship between the number of fishers per 100 sq km of coral reef (participation rate) and the average
annual weight of dry exports per fisher (yield) (P < 0.001; r2 = 0.65; n = 29). The model used log2 transformed data, and
the y-axis (yield) has been back transformed for graphic representation. Thus, for the x-axis, )1 is equivalent to 0.368 (i.e.
e)1) and 8 is equivalent to 2981 (i.e. e8) fishers 100-km)2. Shaded area is the 95% confidence interval. Symbols:
Ac = Coral Sea (Australia), Ag = Great Barrier Reef (Australia), An = Northern Territory (Australia), At = Torres Strait
(Australia), Aw = Western Australia, Cu = Cuba; Eg = Egypt; Er = Eritrea, Fp = French Polynesia, Gp = Galapagos
Islands, Gu = Guam, In = Indonesia, Kb = Kiribati; Ke = Kenya, Ma = Maldives, Mg = Madagascar, Mu = Mauritius,
My = Mayotte, Nc = New Caledonia, Om = Oman, Ph = Philippines, Pn = Papua New Guinea, Sa = Samoa, Se = Sey-
chelles, Sr = Sri Lanka, Ta = Tanzania, To = Tonga, Va = Vanuatu, Wa = Wallis Is.

2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459 45

Management of sea cucumber fisheries S W Purcell et al.
Ecuador, Mexico, Peru, Malaysia, China, Vietnam,
Thailand, Russia), we posit that at least 3 million
people collect sea cucumbers worldwide, either full-
time or alongside other resources.
Resource use, in terms of fishers per unit area,
varied hugely among regions. Participation rates of
export fisheries, in terms of number of fishers km)2
of reef, were highest in Philippines (37.4), Papua
New Guinea (18.1; presently under moratorium)
and Indonesia (15.9). Fishers who collect sea
cucumbers were estimated at 930 000 in the
Philippines, at least 250 000 in Papua New Guinea
and 810 000 in Indonesia. In Indonesia, this
number represents 2.2% of the total estimated
number of fishers in the country (G. Sumiarsa,
unpubl. data). Participation rates in Indian Ocean
and African fisheries averaged 1.7 fishers km)2 of
reef and Pacific Island fisheries had <1 fisher km)2
of reef except for Papua New Guinea (18.1) and
Samoa (22.4). In Samoa, the high participation rate
is sustained through prohibition on exports and
small quantities of sea cucumbers are fished for local
consumption only. The participation rates (fishers
km)2 of coral reef) differed significantly among
tropical fisheries with different status of stocks
(KruskalWallis test; P = 0.044); ranks: moderately
exploited < fully exploited < over-exploited < depleted.
Average landings in tropical fisheries over the
20042008 period were related significantly to
participation rates (Fig. 5; linear model,
F1,27 = 29.34, P < 0.001) but not to coral reef
area (linear model, F1,27 = 0.229; P = 0.636).
Figure 6 Map of the number of regulatory measures employed in global sea cucumber fisheries (n = 76). Symbols ( )
denote fisheries in which moratoria have been set, or already in place, on the collection of all sea cucumber species
for periods of >1 year since 2004.
46
Management measures
The number of regulatory measures varies greatly
among sea cucumber fisheries (Fig. 6). Countries of
Asia and Central and South America usually imposed
far fewer regulations on sea cucumber fishing,
averaging just 1.3 and 2.2 regulatory measures in
fisheries, respectively. Regions with the most regu-
lated fisheries were the North Pacific and North
Atlantic (mean: 6.4 regulatory measures) and indus-
trialized tropical fisheries in Australia (mean: 6.5
regulatory measures). Nonetheless, some fisheries in
other regions have recently had >6 regulatory
measures, for example Cuba, Galapagos Islands,
Japan, Mautitius, eastern Mexico and Papua New
Guinea. Nations in the tropical Pacific and Indian
Oceans averaged 2.5 and 3.1 regulatory measures,
respectively. Although many fishery managers stated
that co-management systems were planned, these
existed in just 12% of sea cucumber fisheries (Fiji,
Galapagos Islands, Kenya, Madagascar, Mexico
(Pacific coast), Niue, Samoa, Saudi Arabia and
Tuvalu).
An nMDS ordination showed that the composi-
tion of regulatory measures used in management
plans varies among regions and fisheries within
regions (Fig. 7). Vectors for most management
measures were associated with more sustainable
fisheries. A high number of species harvested, the
use of moratoria and, to a lesser extent, marine
reserves, were measures associated with over-
exploited and depleted fisheries. 58% of fisheries
No. of regulatory measures
0
1
2
3
4
5
6
7
8
9
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459
 Management of sea cucumber fisheries S W Purcell et al.

2D Stress: 0.15 purpose of their establishment was not specifically

for sea cucumbers. Moratoria on fishing or exports
are in place in 39% of sea cucumber fisheries,
predominantly because of overfishing of stocks
(Fig. 6). Fleet controls (on numbers or sizes or
vessels) exist in just 22% of the fisheries. Of all sea
cucumber fisheries, minimum size limits are im-
posed in 34%, catch quotas (or TACs) in 28% and
gear restrictions in 39%. Rotational harvest strat-
egies (and harvest rules) are practiced in just 5% of
fisheries, all of which are industrialized, and are a
Status relatively new management tool.
Rotatl. harv.
Under expl. Enforc.
Enforcement (and compliance) capacity varied
Mod. expl. Fleet control
Quotas greatly among fisheries and tended to be weak in
Fully expl.
Over expl. Limt. entry tropical fisheries in low-income countries (Fig. 8). In
Licensing
Depleted Size limits some high-income countries, compliance is aided
Gear restr. Seas. cls.
No. reg. meas. with vessel monitoring systems to validate fishing
Moratorium effort in days. In contrast, enforcement and com-
No. Species
pliance in some island countries arises within
co-management systems.
Reserves The PERMANOVA contrasts showed that, in terms of
variables potentially controllable by the fishery
Figure 7 Non-metric multidimensional scaling (nMDS) manager, under-exploited, moderately exploited
plot of variation in the suite of measures controllable by and fully exploited fisheries were similar, but differed
the fishery manager among fisheries in which data on all
significantly from management variables employed
variables were available (n = 67). Fishery points are
in over-exploited and depleted fisheries (Table 2;
distinguished by stock status. Stress = 0.15. Vectors
beneath the plot are derived from Pearson correlations
Fig. 7). The subsequent SIMPER analysis identified
from the normalized data set. that enforcement capacity was the management tool
most strongly related to the dissimilarity between
we assessed had marine reserves. Notwithstanding, unsustainable (over-exploited and depleted; average
many of the reserves are small, weakly enforced rank 1.2) and sustainable (under-exploited, moder-
(McClanahan 1999; Jameson et al. 2002) or in ately exploited and fully exploited; average rank 2.4)
suboptimal habitats for sea cucumbers because the fisheries (Table 2). Nonetheless, other management

Figure 8 Map of enforcement capacity among global sea cucumber fisheries (n = 73). Ranks (1 = low, 2 = moderate,
3 = high) were based on number or compliance officers in each fishery compared to the fishery size and the system
for penalizing infringements.

2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459 47

Management of sea cucumber fisheries S W Purcell et al.

Table 2 Summary statistics of PERMANOVA contrasts and sures important in the majority of difference
SIMPER analyses of variables potentially controllable by between sustainable and unsustainable fisheries (in
the fishery manager across all sea cucumber fisheries for descending order of importance) were the following:
which these data were available (n = 67). Groupings for 1. fleet (vessel) controls,
the SIMPER analysis followed significant differences
2. limited entry controls,
detected in the PERMANOVA contrasts. P -perm are p-values
3. rotational harvest closures,
derived from MCMC permutations.
4. the total number of species harvested by fishers,
5. licensing and reporting requirements, and
PERMANOVA Analysis 6. the number of regulatory measures used by the
manager (Table 2).
Source d.f. Pseudo-F P (perm) Using the same variables as the SIMPER analysis,
a regression tree with four leaves explained 58% of
Overall test across 4 3.38 0.006* the variation in fishery status (Fig. S1). In the
status categories regression tree, the number of species harvested
D vs. O 1 0.98 0.407 explained most of the variance. Similar to the
D vs. [U, M, F] 1 5.22 0.003*
SIMPER analysis, a higher number of species
O vs. [U, M, F] 1 8.69 0.001*
F vs. [U, M] 1 1.12 0.309 harvested was associated with overfished fisheries.
M vs. U 1 0.50 0.736 The next important variable was enforcement
Residual 62 capacity, which corroborates the SIMPER analysis.
Thus, fisheries with more than 4 species fished and
SIMPER analysis a low enforcement capability (32 out of 69 fisheries)
Dissimilarities between have an average status score of 4.22 (equivalent to
groups (U, M and F vs. O and D) over-exploited to depleted). In contrast, if 4 or fewer
species are fished and strong enforcement capability
U, M Dissimilarity
exists (15 of 69 cases), the average status score is
and F O and D contribution
Variable average average (%) 1.73 (under-exploited to moderately exploited).
Fishers in depleted and over-exploited fisheries
harvested three times as many species, on average,
Enforcement 2.4 1.2 9.7
capacity
compared to fishers in better-performing fisheries
Fleet control 50% 5% 8.9 (Table 2; Fig. 7). No-take marine reserves were
Limited entry controls 64% 13% 8.7 employed more often in the over-exploited and
Rotational harvest 14% 0% 8.3 depleted fisheries.
closures
No. species 5.0 15.5 7.9
harvested Human development and sustainability
Licensing and 64% 21% 7.9
Reporting Fisheries in countries with higher HDI tended to
Number regulatory 4.7 2.6 7.7 impose a significantly greater number of regulatory
measures
measures (linear regression; F1,61 = 8.56, P =
Quotas 50% 18% 7.5
Gear restrictions 57% 33% 6.8
0.005). The ANOVA showed that average HDI scores
Seasonal closures 39% 23% 6.8 varied significantly among the five fisheries status
Size limits 50% 31% 6.8 groups (F4,64 = 11.8; P < 0.001) (Fig. 9). Post hoc
No-take reserves 50% 64% 6.6 comparisons identified that average HDI scores were
high (0.890.92) and comparable among under-
D, depleted; O, over-exploited; F, fully exploited; M, moderately
exploited, moderately exploited or fully exploited
exploited; U, under-exploited.
*Significant differences
fisheries but were significantly lower (0.73 and

Averages are on scale of original (un-transformed and stan- 0.63) in over-exploited and depleted fisheries (i.e.
dardized) data. Percentages are percentage of cases in which U = M = F > O = D).
the regulatory measure was used by the management body.

Discussion
variables were also important and contributed only
slightly less to the dissimilarity than enforcement We aimed to relate management measures to
capacity. Specifically, the other management mea- sustainability of sea cucumber fisheries, and our

48 2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459


1.0
Human delopment index ( SE)
0.8
0.6
0.4
0.2
0.0 n =
7 12
d d
te te
loi loi
xp xp
e e -e
der el
y lly
Un ra
t Fu
e
od
M
Figure 9 Mean Human Development Index (HDI) scores
(SE) among fisheries classed within the five different
status categories. HDI scores obtained from United Nations
Development Program data for 2009. Dots display data
points; sample sizes are given at the bottom of each bar.
analyses indicate that resilience of social-ecological
systems (SESs) of sea cucumber fisheries will come
from strengthening enforcement capacity, allowing
only a small number of species to be harvested,
applying input controls, reducing the number of
fishers per unit of fishing ground and improving the
socio-economic state of human communities. An
overarching goal in the management of sea cucum-
ber fisheries should be to safeguard the reproductive
capacity of breeding stocks so that the resources are
available to future generations (FAO 1995, Fried-
man et al. 2008; Purcell 2010). While sea cucum-
ber fisheries are of notable value in coastal artisanal
settings, they are also of traditional value as foods in
some indigenous cultures. Managing exploitation
rates to preserve stocks for subsistence fishing is also
an imperative in maintaining resilience in fishing
communities to external shocks, both economic and
environmental.
Our analysis shows that sea cucumber fisheries
fare worse than most fisheries globally. We found that
58% of sea cucumber fisheries were over-exploited or
depleted, whereas 27% of global stocks (finfish and
invertebrates) were over-exploited or depleted over
the same period (Toral-Granda et al. 2008).
Stocks of commercially valuable species have
been depleted at a comparably fast rate over much
of their distribution (Anderson et al. 2011b). Many
sea cucumber species face a high risk of extinction
through overfishing coupled with inherent biolog-
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459
Management of sea cucumber fisheries S W Purcell et al.
ical and ecological vulnerability (IUCN, in prep.1).
Apart from an unlikely reduction in demand from
Chinese consumers, we believe that sustainability
and resilience of troubled sea cucumber fisheries will
only come from the adoption of radically different
approaches to management.
Enforcement drives sustainability
While marine reserves, international trade agree-
11 25 14 ments and stock assessments may be important
d d ed tools for sustaining sea cucumber stocks (Anderson
te te et
loi loi l
xp xp ep et al. 2011b), our analysis highlights the impor-
r-e D
ve
O tance of enforcement capacity in fisheries sustain-
ability. Indeed, many fisheries struggle with
deterring fishers from illegal activities by compli-
ance measures or strict penalties (Beddington et al.
2007). The SIMPER and RT analyses both indicated
that depleted and over-exploited sea cucumber
fisheries predominantly had weak enforcement
capacity. Thus, fine-tuning management regula-
tions or developing complex management plans are
less likely to succeed in protecting minimum viable
populations of sea cucumbers than investment in
compliance and enforcement.
In Papua New Guinea, a national management
plan in 2001 included minimum size limits for dried
and fresh animals, prohibition on compressed-air
diving and torches, seasonal closures, no-take
marine reserves, a global TAC and limited entry to
PNG citizens (Kinch et al. 2008b). However, mon-
itoring and enforcement costs and the sheer number
of fishers made it impossible to curtail illegal fishing
practices with a centralized management system;
stocks of nearly all commercial species declined by
more than 80%, prompting a national moratorium
in 2009 (Purcell 2010). Moratoria may be effective
at saving stocks from extirpation because catches
can be monitored generally at the point of export.
However, illegal trade may still need wider enforce-
ment. In Panama, for example, the government
issued a Ministerial Decree prohibiting the collec-
tion, processing or sale of sea cucumbers but illegal
fishing still occurs on both coasts (R. Pretto,
personal communication). The case-study from
Papua New Guinea (Kinch et al. 2008b) leads us
to agree that management plans must be developed
1
IUCN Red List of Threatened Species: Assessment of species within
the order Aspidochirotida. In review. Also see Polidoro et al.
(2011).
49
Management of sea cucumber fisheries S W Purcell et al.
more with stakeholders, namely fishers and proces-
sors, but predicated on the enforcement capacity.
Top-down enforcement in small-scale fisheries
with numerous and widely dispersed fishers is
intractably difficult so more participatory ap-
proaches are needed (Berkes et al. 2001; Cochrane
and Doulman 2005; Orensanz et al. 2005; Bene et al.
2007). We stress that compliance in sea cucumber
fisheries will be best achieved through enforcement
at the narrow middle of the hourglass of the trade
chain within each fishery. Forcing compliance of
minimum sizes and permissible species at centralized
processing agents and export hubs should feedback
to compliance by fishers (Purcell 2010).
Management measures
Resource managers have been recently called upon
to apply a diverse set of essential regulatory
measures (Garcia et al. 2003; Stefansson and
Rosenberg 2005; Worm et al. 2009). The depleted
and over-exploited fisheries we examined tended to
have far fewer regulatory measures (mean: 2.6)
than better-performing fisheries (mean: 4.7).
Although the comparisons have some inherent bias
(e.g. young temperate fisheries with, as yet, healthy
stocks tend to have many regulations), the findings
indicate that a set of at least 4 or 5 key regulatory
measures is needed in sea cucumber fisheries
(Table 2; Fig. 7). Based on our SIMPER analysis,
the most successful regulatory measures appeared
to be limited entry controls (elaborated later), fleet
(capacity) controls, restrictions on the number of
harvested species and licensing requirements.
Applying further measures should depend upon
the technical capacity of the management agency
and idiosyncrasies of the fishery (Purcell 2010). A
set of regulations is needed (FAO 2010b, Purcell
2010), but the management plan should not
contain so many regulatory measures that compli-
ance and enforcement become so difficult as to be
practically impossible (Cochrane 2002).
In fisheries generally, tighter capacity limits are
needed on the number and size of vessels (FAO
1995, Pauly et al. 2002) and should apply to sea
cucumber fisheries (Purcell et al. 2009a). Restrict-
ing the size of boats in the fishery probably has
several advantages. Small boats cannot easily ven-
ture to offshore sites, thus limiting sequential
exploitation (Berkes et al. 2006) and allowing
offshore sanctuaries to potentially subsidize larval
supply to inshore fishing grounds. Larger boats also
50
permit multiday trips that tend to promote excessive
harvests (Purcell 2010). Controls on boat size can
also avoid overcapitalization that can drive catches
beyond small scale to meet capital rent. States are
urged to protect the rights of small-scale and
subsistence fishers to meet their livelihoods needs
(FAO 1995, Bene et al. 2007), which can be
achieved in part through limits on industrialized
fishing.
As illustrated by SIMPER and RT analyses, sea
cucumber fisheries exploiting more species tended
towards overfishing; fishers in over-exploited and
depleted fisheries collected three times as many sea
cucumber species as those in better-performing
fisheries. Apparently, stocks of medium- and high-
value species are exposed to greater risk of overf-
ishing in fisheries where fishers shift to ever broader
range of lower-value species (Kinch et al. 2008a;
Friedman et al. 2011). Allowing fishers to exploit a
broad range of species opens the door to opportu-
nistic depletion, whereby high-value species are
collected despite low abundance because lower-
value species can be fished in the same habitat and
using the same methods. Managers should take
steps to conserve fishery ecosystems (FAO 1995).
Relatively small and adaptive lists of species (e.g.
medium and high-value) permitted to be caught and
sold (Purcell et al. 2009a) should thus act to
mitigate opportunistic depletion and to preserve
the role of some populations in sediment health and
nutrient recycling (discussed earlier). Regression
tree analysis indicated that most fisheries targeting
more than 4 to 5 were usually over-exploited or
depleted.
Poverty, participation rates and fisheries
sustainability
Sea cucumber fisheries in low-HDI (low-income)
countries are relied upon heavily for poverty
reduction and/or subsistence diets (Choo 2008a;
Conand 2008; Kinch et al. 2008a,b, Toral-Granda
2008b). Tragically, our data show that these tended
to experience overfishing significantly more than
fisheries in high-HDI (high-income) countries. Cin-
ner et al. (2009) found that fish stocks in the
western Indian Ocean were affected by socio-
economic development in a U-shaped relationship,
whereby over-exploitation appears highest at inter-
mediate levels of human community development.
In contrast, our findings of increasing depletion of
sea cucumber stocks with lower levels of human
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459

development show that the U-shaped relationship
does not hold for some other marine resources. We
posit that the biological and ecological vulnerability
of sea cucumbers and the ease in which they can be
collected allows overfishing of stocks of even the
very poor and under-developed societies.
Our results support that low-HDI countries have
a much harder time controlling overfishing and that
poverty drives poor fishers to deplete stocks to levels
far below those at which bioeconomic equilibrium
(see Beddington et al. 2007) would normally be
met. Sea cucumber fisheries in high-HDI countries
are often more recent or in a developmental phase
and those fishers often have social welfare or other
livelihood alternatives (Purcell 2010). Part of the
upsurge of sea cucumber fisheries in developed
countries appears to be driven by strengthening
markets as supply from depleted fisheries in devel-
oping countries wanes. We draw two important
conclusions: (1) that support for improved fisheries
management and enforcement is needed over-
whelmingly in low-income countries, and (2) that
sustainability of sea cucumber fisheries in low-HDI
countries must come from holistic aid that provides
fishers with other livelihood options and improves
the desperate social context in which resources are
exploited (Andrew et al. 2007; Bene et al. 2007).
Fisheries in low-HDI countries also tend to have
much higher participation rates of fishers. For
instance, whereas there are more than one and a
half million sea cucumber fishers in Indonesia and
Philippines combined, there are <100 sea cucumber
fishers in Australia (Kinch et al. 2008a) and in
Canada (Hamel and Mercier 2008a) and about one
hundred in the Seychelles (Aumeeruddy and
Conand 2008). Our analysis indicates a range of
about 2 to 10 fishers 100-km)2 of tropical reef
where after yields per fisher become predictably
paltry and the race-for-fish drives exploitation to
unsustainable levels (see Fig. 5). Conversely, partic-
ipation rates below about 1 fisher 100-km)2 should
provide high annual yields. Some of the uncertainty
in yields at lower participation rates relates to
productivity of reef systems. Wallis Island, for
example, had relatively low participation rates com-
pared to available habitat but the low productivity
and larval connectivity of such isolated reefs prob-
ably limits potential yields to fishers (Kinch et al.
2008a). In some fisheries, like the Philippines, it will
be near impossible to regulate the number of fishers
in the current open-access system afflicting the sea
cucumber fisheries. Thus, other measures will need
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459
Management of sea cucumber fisheries S W Purcell et al.
to be relied upon. Where possible, managers should
limit the number of fishers and our findings should
serve as a valuable guide to a sensible number of
fishers knowing the reef area of the fishery.
Measures to mitigate loss of biodiversity
No-take reserves were common within the sea
cucumber fisheries we examined, and some fisheries
relied on reserves as the sole management tool.
Surprisingly, we found no correspondence between
the presence of reserves and the sustainability of
fisheries. In fact, reserves were more common in the
depleted and over-exploited fisheries (see Table 2).
Despite a trend in developing countries for large
MPAs, they are still generally small in most less-
developed countries and therefore more easily allow
illegal fishing activities (Sale et al. 2005; Purcell
2010).
Few reserves globally are designed for sea cucum-
bers and few, if any, appear to have been added as
reactive measures after declines of these resources.
However, our data do not refute the potential for
no-take reserves to act as safeguards to the loss
of egg production in the fishery (Browman and
Stergiou 2004; Sale et al. 2005). But if compliance
of reserve boundaries is weak, poaching may negate
any such benefits (Sethi and Hilborn 2008). For
example, there are more than 150 marine reserves
in the Philippines but many are small and/or with
weak enforcement so their effect on recruitment
subsidization is questionable (Choo 2008b). Some
reserve sites may not favour the development of
dense sea cucumber populations (Purcell et al.
2009a), perhaps owing to food availability, appro-
priate shelter, settlement rates or post-settlement
processes. Thus, we conclude that marine reserves
representing just a small proportion of habitats will
not suffice as a sole management measure to
safeguard resilience in sea cucumber fisheries. Sites
with ideal habitats or populations of sea cucumbers
may also need to be selected specifically in a
network of reserves. Although not designed specif-
ically for protecting sea cucumbers, no-take zones
on the Great Barrier Reef, Australia, have 34 times
higher densities of the high-value species
Holothuria whitmaei (Uthicke and Benzie 2000).
Populations on fished reefs probably are not effec-
tively reproducing at low densities (Uthicke 2004),
and their recovery may now depend on the supply
of larvae produced elsewhere (e.g. from the no-take
zones).
51
Management of sea cucumber fisheries S W Purcell et al.

Most of the commercially important sea cucum-
ber species are widely distributed and harvested in
multiple countries. Over-exploited or severely
depleted fisheries may thus benefit from interna-
tional agreements to regulate trade (e.g., CITES).
The IUCN recently evaluated over 300 aspidochi-
rotid sea cucumbers for inclusion on the Threatened
Species Red List, categories pending final review
(Polidoro et al. 2011). Based on significant (>80%)
declines in populations over the majority of their
range over the past three generations, a handful of
tropical species will likely be listed as Endangered and
warrant further listing in CITES. Currently, only one
species (Isostichopus fuscus) is listed in CITES within
Appendix III (Altamirano et al. 2004). Impediments
to CITES listing of other species include limited
biological studies, uncertainty about population
status, difficulties in distinguishing processed (dried)
animals in trade and concerns that CITES
listing may spur black-market trade (Choo
2008a). While the IUCN Red List is not a
binding agreement, it may provide impetus for
CITES listing and for resource managers to take
independent steps to improve the conservation
of threatened species.
Pulse vs. continuous fishing
Our review demonstrated variation and uncertainty
in the resilience of sea cucumber stocks. Rapid
decimation of stocks within 24 years of fishing has
been recorded in Egypt (Lawrence et al. 2004;
Hasan 2005), Oman (Al-Rashdi and Claereboudt
2010), and Tonga (P. Ngaluafe, unpubl. data).
While Friedman et al. (2011) advocate a strategy of
closure-pulse fishing-closure, we caution that the
social and ecological risks must be considered.
Obliging fishers and processors to shift to other
income streams in years of closure may be socially
unacceptable. As has recently occurred in Tonga,
this strategy may also spur public pressure to
prolong pulse fishing events beyond safe ecological
limits (P. Ngaluafe, personal communication). Sup-
ply-chain arrangements would also be dislocated in
years of closure. In contrast, several decades of
modest continuous fishing within a rotational

(a)
Stock abundance

Pulse fishing
Continuous fishing
(modest rate)

Allee threshold

Time (years)

(b)
Stock abundance

Continuous fishing
(modest rate)

Allee threshold
Pulse fishing

Time (years)

Figure 10 Hypothetical variations in stock abundance under two different harvest strategies: pulse fishing; modest
continuous fishing. Fishing periods are denoted by dashed curves; periods of fishing closures are denoted by solid curves.
Under a closure-pulse fishing-closure strategy, populations are reduced at a faster rate than under a modest continuous
fishing strategy and risk falling below levels at which depensatory (Allee) effects occur. (a) The Allee threshold is low
and stock abundance is not suppressed below it during the intensive pulse fishing and stocks recover thereafter. (b) The
Allee threshold is slightly higher causing stock abundance after the second pulse fishing period to be below the level at
which per capita growth of the population is positive, and the population fails to recover thereafter.

52 2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459


harvest strategy has resulted in apparently stable
stocks in British Columbia (Canada).
In addition, short and intense pulse fishing events
may lower stocks beyond thresholds at which
depensatory effects erode the ability of stocks to
replenish into the future (Fig. 10). While repeated
surveys would be needed to monitor the progression
of pulse fishing effects on stocks, there is often limited
capacity for managers to collect and analyse such
data (Friedman et al. 2011). Moreover, the signals of
stock depletion and management responses could be
too late to curtail fishing so that populations remain
above reproductively effective levels. We argue that
continuous (non-disjunct) fishing at modest rate
would serve better at maintaining effective breeding
densities of sea cucumbers than a closure-pulse
fishing-closure strategy (Fig. 10).
For the high-value Pacific black teatfish, H. whit-
maei, annual catches of <5% of virgin biomass over
1015 years were shown to reduce breeding popu-
lations by 75 per cent on the Great Barrier Reef
(Uthicke 2004). Therefore, modest fishing to avoid
over-exploitation would need to be in the range of just
24% of unfished biomass per year. In the absence of
reliable estimates of fishable biomass of particular
species, catch rates must be curtailed very conserva-
tively. Risk in a pulse fishing strategy arises because
thresholds of population abundance for Allee effects
(depensation) are uncertain for sea cucumbers (Bell
et al. 2008) and intensive overfishing can then lead
to negative population growth if abundances drop
too low before fishing closures are enforced.
Conclusions
Sea cucumbers are low-food-chain organisms that
have most recently succumbed to pandemic overf-
ishing. There is increasing evidence that they bolster
ecosystem health and contribute to the resilience of
coral reef, soft-bottom and deep-water habitats.
Much of the fishing of tropical sea cucumbers
occurs on coral reefs and lagoons, which are under
particular threat from global impacts such as
climate change and ocean acidification (Hoegh-
Guldberg et al. 2007; Fabricius et al. 2011). Provid-
ing high resilience to these habitats by managing
the exploitation of key taxa is a possible way to
rescue these reefs or at least buy time until global
solutions have been found (Marshall and Johnson
2007). However, our review indicates that few sea
cucumber fisheries currently set, or achieve, this
ecosystem goal.
2011 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 14, 3459
Management of sea cucumber fisheries S W Purcell et al.
Recent assertions about improved management of
sea cucumber fisheries have centred on regulatory
measures and management actions (FAO 2010a,
Purcell 2010), marine reserves and international
regulations (Anderson et al. 2011b) research, mon-
itoring and harvest strategies (Friedman et al.
2011). Our findings reveal that a new paradigm is
needed for managing sea cucumber fisheries; one in
which resources are shifted from the development of
complex management plans to enforcement and
compliance of simple sets of regulations and to
tackling the socio-economic challenges of coastal
fishers that transcend fisheries. The best devised
management plans will fail if disincentives to illegal
fishing activities are not strong enough (Beddington
et al. 2007) and/or if underlying poverty of fishers is
not improved (Andrew et al. 2007; Cinner et al.
2009; Allison et al. 2011).
We also conclude that multiple management
measures are needed in sea cucumber fisheries (FAO
2010a, Purcell 2010) but not so many that they
cannot be easily understood and enforced. In
addition to compliance and enforcement, the key
regulatory measures appeared to be the following:
1. a small list of permissible species for exploitation,
2. fleet controls, especially on the size of boats in
the fishery,
3. limited entry controls to restrict the number of
fishers, and
4. licensing and reporting requirements.
Recent evidence of slow growth, low natural
mortality and high longevity in some species (Hamel
and Mercier 1996; Uthicke et al. 2004; So et al.
2010) underscores the need for more conservative
management strategies than in the past and cautions
the use of conventional fisheries science underpinned
by estimations of maximum sustainable yield. The
great variation in the scale of fishing activities,
management systems and technical capacities of
management bodies means that multiple, country-
or region-specific, solutions will be needed to redress
the shortcomings in collapsed fisheries. Operational-
izing an ecosystem approach to fisheries manage-
ment (sensu Garcia et al. 2003) requires a greater
involvement of stakeholders, consideration of alter-
native management systems and a higher priority of
social science in management institutions. Resilience
in sea cucumber fisheries will come about only if
fishers are part of the management system and can
adapt quickly to changes in the resource (Lebel et al.
53
Management of sea cucumber fisheries S W Purcell et al.
2006; Andrew et al. 2007). In the vast majority of
global fisheries, regulations towards sustainability
are commonly undermined by political pressures
(Mora et al. 2009). Transformation of the manage-
ment paradigms currently undermining sea cucum-
ber stocks, therefore, inevitably needs to be supported
by decision-makers resolutely if these animals are to
remain valuable to the livelihoods of coastal peoples
and provide the eco-services that contribute to
healthy marine ecosystems.
Acknowledgements
For the provision of fishery data and information, we
graciously thank H. Eriksson (Zanzibar, Tanzania,
Mayotte), T. Lavitra (Madagascar), C. Macamo, L.
Muaves and S. Fernando (Mozambique), H. Abdere-
mane and A. Stroud (Comoros), Y. Robinson (Sey-
chelles), C. Dissanayake (Sri Lanka), K.
Tesfamichael, S. Mahmud, R. Dabbagh and B. e-
Lengeh (Iran), N. Cariglia, B. Allison and S. Hassan
(Maldives), M. Slater (New Zealand and Tanzania),
S. Kundun, S. Jhummun and S. Meunier (Mauritius),
M. Trianni (CNMI), A. Stein (French Polynesia), B.
Ponia and K. Raumea (Cook Islands), T. Skewes
(Torres Strait, Western Australia, Great Barrier
Reef), R. Donald (Vanuatu), L. Gisawa and J. Kinch
(Papua New Guinea), J. Samuelu (Samoa),
P. Ngaluafe (Tonga), B. Mugneret (Wallis and
Futuna), D. Nguyen (Vietnam), C. Ramofafia and
R. Maneiria (Solomon islands), M. Depaune (Nauru),
L. Labe and R. Gamboa (Philippines), G.Sumiarsa
(Indonesia), B. Tibbatts (Guam), I. Eeckhaut (Oman),
N. Duprey (British Columbia), E. Hynick and D.
Stansbury (Newfoundland and Labrador), P. Kalvass
(California), Robert Watts (Maine), S. Rowe and C.
MacDonald (Nova Scotia), B. Meredith (Alaska), G.
Thorarinsdottir (Iceland), S. Luna and P. Martinez
(Ecuador), G. Perez, E. Gomez and M. Benavides
(Colombia), J. Alvarado (Costa Rica), F. Solis-Marn
(Mexico), C. Ibanez and C. Guisado (Chile), J. Palazon
Fernandez, C. Hasbun and E. Barraza (El Salvador),
B. Brenes (Nicaragua), Y. Hooker (Peru), N. Serrany
(Panama) and F. Scarabino (Uruguay). We are
grateful to F. Carocci for graphic work on the
figures. This study was supported by the National
Marine Science Centre, Southern Cross University
and the Pacific Agribusiness Research for Develop-
ment Initiative (PARDI) of the Australian Centre for
International Agricultural Research (ACIAR)
(SWP), the World Wildlife Fund (MVT-G) and the
Food and Agriculture Organization of the United
54
Nations (AL). We are grateful to two anonymous
reviewers for their constructive comments, which
helped improve the manuscript.
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Supporting Information
Additional Supporting Information may be found in
the online version of this article:
Figure S1. Regression tree of the measures
controllable by the fishery manager that best
explained the relationship with different status
ranks of the fisheries (15 for under-exploited to
depleted status). Status values for the different cut-
offs are given at the bottom of the tree, with sample
sizes below, of fisheries falling into each group.
Table S1. (a) Status and general characteristics
of the various fisheries examined. (b) Regulatory
measures useda in the various fisheries examined.
(c) Measures used (continued)a, participation, yields
and information sources for the various fisheries
examined.
Please note: Wiley-Blackwell are not responsible
for the content or functionality of any supporting
materials supplied by the authors. Any queries
(other than missing material) should be directed to
the corresponding author for the article.
59