F I S H and F I S H E R I E S , 2016, 17, 176–192

Co-management in Latin American small-scale shellfisheries:

assessment from long-term case studies

Omar Defeo1,2, Mauricio Castrej
on3,4, Roberto P
erez-Casta~

neda5, Juan C Castilla6, Nicol
as L Guti
errez7, Timothy
8 9,10
E Essington & Carl Folke

1
UNDECIMAR, Facultad de Ciencias, Igu
a 4225, 11400, Montevideo, Uruguay; 2GEPEIA, Centro Universitario de la
on Este, Ruta nacional N°9 intersección con Ruta N°15, Rocha, Uruguay; 3Interdisciplinary PhD program,
Regi

Dalhousie University, Halifax, Nova Scotia, B3H 4R2, Canada; 4World Wildlife Fund, Calle Piqueros s/n y 13 de
Febrero, Puerto Ayora, Santa Cruz, Galapagos, Ecuador; 5Facultad de Medicina Veterinaria y Zootecnia, Universidad
onoma de Tamaulipas, A.P. 263, Cd. Victoria, 87000, Tamaulipas, M
exico; 6Centro de Conservaci

Aut
on Marina
(N

ucleo Milenio) & Estaci

on Costera de Investigaciones Marinas, Las Cruces, Facultad de Ciencias Biol

ogicas, P.
olica de Chile, Casilla 114-D, Santiago, Chile; 7Marine Stewardship Council, 1 Snow Hill, London, EC1A
Universidad Cat

2DH, UK; 8School of Aquatic and Fishery Sciences, University of Washington, Seattle, WA, 98195, USA; 9Stockholm
10
Resilience Centre, Stockholm University, SE - 106 91, Stockholm, Sweden; Beijer Institute, Royal Swedish Academy of
Sciences, SE - 104 05, Stockholm, Sweden

Abstract Correspondence:
Omar Defeo
Co-management (Co-M), defined as the sharing of management tasks and responsi-
UNDECIMAR
bilities between governments and local users, is emerging as a powerful institu- Facultad de Ciencias
tional arrangement to redress fisheries paradigm failures, yet long-term Igu
a 4225
assessments of its performance are lacking. A comparative analysis of five small- 11400 Montevideo
scale Latin American shellfisheries was conducted to identify factors suggesting Uruguay
success and failure. In Chile, Uruguay and Mexico Co-M produced positive effects, Tel.: 5982-5258618
Fax: 5982-5258617
including stabilization of landings at low levels, increase in abundance, CPUE, unit E-mail: odefeo@
prices and revenues per unit of effort, and reduced interannual variability in sev- dinara.gub.uy
eral fishery indicators, particularly in landings. Co-M was successful because it was
mainly bottom-up implemented and accompanied by-catch shares (spatial property
Received 5 Apr
rights and community quotas). By contrast, Co-M implementation was unable to 2014
prevent the collapse of the Galapagos sea cucumber fishery, as reflected by a Accepted 19 Sep
decrease in abundance and CPUE. Negative effects were also observed in the Gala- 2014
pagos spiny lobster fishery during Co-M implementation. However, recovery was
observed in recent years, reflected in a stabilization of fishing effort and the highest
CPUE and economic revenues observed since the beginning of the Co-M implemen-
tation phase. The combined effects of market forces, climate variability and a mora-
torium on fishing effort were critical in fishery recovery. We conclude that Co-M is
not a blueprint that can be applied to all shellfisheries to enhance their governabil-
ity. These social–ecological systems need to be managed by jointly addressing prob-
lems related to the resources, their marine environment and the people targeting
them, accounting for their socioeconomic and cultural contexts.

Keywords Co-management, fishery indicators, Latin America, small-scale

shellfisheries, social–ecological systems

176 DOI: 10.1111/faf.12101 © 2014 John Wiley & Sons Ltd

 Co-management in small-scale fisheries O Defeo et al.

Introduction 177
Methods 178
Chile: Concholepas concholepas, ‘loco’ 178
Uruguay: the yellow clam Mesodesma mactroides 179
Galapagos Islands (Ecuador): the sea cucumber Isostichopus fuscus and spiny lobsters 180
Panulirus penicillatus and P. gracilis
Mexico: the spiny lobster Panulirus argus in Yucatan Peninsula 181
Analysis 181
Results 181
Discussion 186
Conclusions 189
Acknowledgements 190
References 190
Supporting Information 192

world’s capture fishers (Kalikoski and Franz

Introduction
2014). In these fisheries, novel adaptive gover-
Fisheries are complex social–ecological systems nance, notably the adoption of community-based
(Berkes and Folke 1998) in crisis. The world fishery co-management (hereafter co-management: Co-M)
crisis is not a single issue but a series of complex, arrangements, has been suggested as a useful step
interdependent and multifaceted challenges of differ- towards sustainability (Castilla and Defeo 2001;
ent types of fisheries, communities, societies and McClanahan et al. 2009; Evans et al. 2011). Co-
markets, which require different strategies and tools M, defined as the sharing of responsibilities
for problem-solving (Castilla and Defeo 2005; Hil- between governments and local users, is emerging
born et al. 2005; Ostrom 2007, 2009; Worm et al. as a powerful governance mode to redress fisheries
2009). For example, the marked differences in the paradigm failures (Carlsson and Berkes 2005;
intrinsic properties of large-scale industrial fisheries Guti
errez et al. 2011; Cinner et al. 2012). Advan-
and small-scale coastal artisanal fisheries lead to tages of Co-M include the following (Berkes et al.
important differences in the potential effectiveness of 2001; Castilla and Defeo 2001; Basurto and
management measures. In industrial fisheries, the Ostrom 2009; Schultz et al. 2011): (i) enhance-
availability of large amounts of detailed data, techni- ment of ownership and fishing rights among fish-
cal expertise and governmental funding facilitates ers, encouraging more responsible harvesting; (ii)
relative precise assessment of status and the adop- encouragement of collective ownership and com-
tion of classic management tools. These tools pliance by including fishers in the decision-making
include catch quotas, effort restrictions and techni- process; (iii) a more sensitive policy and manage-
cal gear modifications to promote long-term social, ment to local knowledge and socioeconomic and
ecological and management objectives (Caddy ecological restraints; (iv) greater incentives for reli-
1999; Hilborn 2007; Hilborn and Hilborn 2012). able monitoring, control and surveillance by fish-
By contrast, most small-scale fisheries lack the cen- ers themselves; and (v) increased capability of
tralized enforcement capacity and data collection fishers to engage in adaptive management with
needed to ensure compliance with regulations and continuous learning as a central part of the co-
to diagnose biologic and socioeconomic status. This management process. Hence, Co-M demands
makes traditional stock assessments difficult or intensive fishers’ participation through the incor-
impossible to conduct, precluding the management poration of their scrutiny and responsibility in
of small-scale fisheries on the basis of catch quotas governance schemes (Chuenpagdee and Jentoft
and effort control. 2007).
Small-scale coastal fisheries, mostly occurring in Despite the increasingly widespread adoption of
developing countries, account for ca. 40% of world Co-M for solving governance issues in coastal
fish catches (ca. 90 million t, FAO 2014) and pro- social–ecological systems (Cinner et al. 2012),
vide direct employment for >90% out of 39 million long-term assessments of its success in terms of

© 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192 177
Co-management in small-scale fisheries O Defeo et al.

ecological–socialeconomic performance indicators factors associated to their success or failure.

are few (Ostrom 2009). Most human and eco- Finally, we discuss future needs, challenges and
nomic resources from governments and external issues to improve shellfisheries Co-M.
donors have been allocated to develop and imple-
ment Co-M as a novel institutional arrangement,
Methods
whereas very limited or no exertion has been allo-
cated to monitor and periodically assess this gover- The shellfisheries analysed here include the muricid
nance mode. Guti
errez et al. (2011) examined 130 gastropod Concholepas concholepas (Muricidae),
co-managed fisheries in a wide range of countries ‘loco’, in Chile; the yellow clam Mesodesma mactro-
with different degrees of development, ecosystems, ides (Mesodesmatidae) at Barra del Chuy, Uruguay;
fishing sectors and type of resources, and high- the sea cucumber Isostichopus fuscus (Stichopodidae)
lighted the lack of long-term databases for testing and spiny lobsters Panulirus penicillatus and P. graci-
specific hypotheses about Co-M performance. They lis (Palinuridae) in the Galapagos Islands, Ecuador;
showed that a low number of cases included a full and the spiny lobster Panulirus argus (Palinuridae)
evaluation of Co-M failure or success: only 6% pre- in the Yucatan Peninsula, Mexico. The selection of
sented before–after, control–impact, or complete these case studies was not at random, because the
before–after control–impact (BACI) approaches. lack of suitable data for most fisheries makes it
These studies are critical to produce well-grounded impossible to perform a randomized selection of Co-
evidence about the conditions that facilitate or M shellfisheries in Latin America. However, this
preclude successful Co-M arrangements. study includes the most comprehensive long-term
Most small-scale shellfisheries in Latin America shellfisheries datasets compiled in this region to
are based on high-value species that are vital to date, including a full range of social, ecological, cul-
the livelihoods of coastal fishing communities (De- tural and governance settings (Table 1).
feo and Castilla 2005, 2012; Orensanz et al.
2005, 2013). The commercial importance of these
Chile: Concholepas concholepas, ‘loco’
social–ecological systems has gradually increased
as finfish fisheries have either reached full exploi- Concholepas concholepas is a unique muricid gastro-
tation or experienced over-exploitation (Castilla pod restricted to rocky shores of Chile and south-
and Defeo 2001). In Latin America, coastal shell- ern Peru. Economically, it is one of the most
fisheries have been mostly developed along important benthic shellfish mollusc extracted in
extended coasts under open access regimes. This Chile by more than ca. 22 000 ‘hookah’ profes-
has made their governance extremely challenging sional divers (Table 1) and an unknown number
for two main reasons: (i) the number of extractors, of snorkelled skin-divers and food gatherers
either authorized or unauthorized (e.g. free riders, (Dur
an et al. 1987). The loco was over-exploited
recreational users) and landing sites cannot be in the late 1980s, leading to a 4-year full fishery
readily controlled; and (ii) operational manage- closure (1989–92). In 1991, the Fishery and
ment measures are almost impossible to enforce Aquaculture Law No. 18 892 (FAL) drastically
and are beyond the finances of most management reformed the access to fishery resources along the
agencies. Thus, after many decades of intensive coast of Chile. The FAL differentiated artisanal and
fisheries extraction, exacerbated by coastal degra- industrial subsectors and introduced coastal sea-
dation and globalization of markets, many shellfish zoning schemes (Castilla 2010). Moreover, the
are near the point of functional extinction (Car- 1991 FAL incorporated, among other manage-
ranza et al. 2009; Beck et al. 2011). In these shell- ment tools, national and regional fishery closures,
fisheries, Co-M is emerging as a promising the allocation of community quotas for fully
governance mode to solve the fishery crisis, exploited resources and territorial user rights for
through the active involvement of local communi- fisheries (TURFs), known as management and
ties (Basurto 2005; Defeo and Castilla 2005; Hil- exploitation areas for benthic resources (MEABRs),
born et al. 2005; Basurto and Ostrom 2009; to artisanal fisher associations. The national confed-
Begossi et al. 2011; Silvano et al. 2014). In this eration of artisanal fishers, which groups all arti-
study, we assess the long-term performance of Co- sanal fisher associations, had a primary role during
M in five Latin American shellfisheries using the preparation and implementation of this major
multiple fishery indicators and identify the main governance shift. Since the effective implementation

178 © 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192
 Co-management in small-scale fisheries O Defeo et al.

Table 1 Summary information of the five small-scale Latin American shellfisheries analysed in this paper.

Loco Yellow clam Spiny lobster Sea cucumber Spiny lobster

Country Chile Uruguay Ecuador Ecuador Mexico

Region/locality Central/El Quisco Rocha/Barra Galapagos Galapagos Quintana Roo/Pta.
del Chuy Allen
Habitat type Subtidal, rocky Intertidal, Subtidal, rocky Subtidal, rocky Subtidal, soft-bottom
sandy beach
Number of active ca. 22 000 40 410 536 76
fishers
Economic value Up to 50 0.24 0.69 2.78 1.00
(million US$)
Main market Export Domestic Export Export Domestic
(Domestic/Export)
Individual revenues 280 fisher
1 h
1 21 fisher
1 h
1 51 fisher
1 h
1 110 fisher
1 h
1 366 boat
1 d
1
(US$) per unit
of effort1
Fishing method Hand-gathering by Hand-gathering Hand-gathering by Hand-gathering by Hand-gathering
hookah and skin-divers (shovels) hookah and hookah and by skin-divers
skin-divers skin-divers
Organization Unions Family groups Cooperatives Cooperatives Cooperatives
Year of Co-M 1993 1988–90 1997–2002 1998–2002 1969
implementation
Methodological
approach
Control–impact Comparison between Comparison between
open access (OA) weakly managed
and Co-M sites2 and Co-M sites
(1998–2002)
Before3 1981–88 1982–87 1997–2002 1999–2002 1986–2003
After 1994–2009 1990–94 2003–12 2003–13

1
Revenue per unit of effort is given for fishing seasons that varied from days to months.
2
Information extracted from previous papers and unpublished information (J. C. Castilla).
3
Missing years of data between before and after periods are due to fishery closures.

of the FAL in 1993, more than 700 MEABRs have included catch, fishing effort, CPUE and unit price
been allocated exclusively to small-scale fisher asso- paid for loco at the Caleta. The information gath-
ciations through non-transferable user rights (Ca- ered allowed the comparison of fishery indicators
stilla 2010; Gelcich et al. 2010). Accountability before and after Co-M implementation at national
systems in place at each MEABR include stock and local scales (Table 1).
assessments and total allowable catch (TAC) alloca-
tions decreed by the government (Gelcich et al.
Uruguay: the yellow clam Mesodesma mactroides
2010). MEABRs are the only legal extractive fishing
grounds of loco in the country. Mesodesma mactroides is a sedentary infaunal
Our assessment for the loco fishery covered dif- bivalve artisanally exploited (shovels and hand-
ferent time periods and scales. At national level, picking) in sandy beaches of Brazil, Uruguay and
official landings from 1981 to 2009 (Table 1) Argentina. In Uruguay (Table 1), the yellow clam
were obtained from the Servicio Nacional de Pesca is mainly found along 22 km of sandy beach
(SERNAP). Unit export prices were obtained by the (Barra del Chuy). This place represents the only
ratio between economic revenues and exportation one in the country where the species is exploited.
volumes obtained from the Servicio Nacional de Catches were low before the 1980s, increasing up
Aduanas (Chile). Information at a single Caleta (El to 3.5 times between 1981 and 1985 (up to
Quisco) was gathered in 1988 (before Co-M) and 250 t) and then decreasing more than 50% in
from 1995 to 2002 (after Co-M). Fishing metrics 1986. Afterwards, coastal marine authorities, sci-

© 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192 179
Co-management in small-scale fisheries O Defeo et al.

entists and local fishers agreed on the implementa- divers in subtidal rocky habitats (Table 1). At
tion of a full fishery closure, which extended from the mid-1990s, management of marine resources
April 1987 to November 1989 (Defeo 1996). Two faced several political and socioeconomic chal-
years after the closure, adult clam density lenges, including the over-capitalization of the
increased by more than 400%, and thus, the fish- local small-scale fleet as result of the unregulated
ery was reopened from December 1989 onwards, expansion of the sea cucumber fishery (Castrej
on
with the implementation of several management 2011). Increasing social conflicts and ecological
strategies (Defeo 1996): (i) allocation of a degradation led to the declaration of the Galapagos
restricted number of fishing licenses to those fish- Special Law (GSL) in March 1998 and to the crea-
ers with longer activity in the fishery (the number tion the Galapagos Marine Reserve (GMR), a mul-
of fishers was estimated as the ratio between the tiple-use reserve of nearly 138 000 km2. Several
net economic value of the overall quota and two management measures were gradually imple-
times the minimum national wage decreed by the mented to shift from an open access to a common
government); (ii) establishment of a monthly TAC property regime in fishery resources between 1998
and an individual quota per fisher; (iii) design of a and 2002, including the following: (i) extension of
spatial management scheme featured by an infor- the boundaries of the reserve from 28 to 74 km;
mal (i.e. not legally recognized) allocation of (ii) prohibition of industrial fishing; (iii) establish-
co-ownership authority to groups of fishers in ment of a moratorium on new entrants to fisher-
beach management units with well-defined ies; (iv) allocation of licenses and fishing permits
boundaries. This encouraged an active voluntary exclusively to local artisanal fishers; and (v) an
fisher participation in enforcing regulations, thus unsuccessful attempt to implement a catch share
generating a de facto Co-M regime (Castilla and programme in the sea cucumber fishery in 2001.
Defeo 2001). The Co-M phase of the fishery However, two of the most relevant measures were
lasted until late 1994, when mass mortalities deci- the institutional shift from a hierarchical (top-
mated populations of M. mactroides throughout its down) to a Co-M regime, and the implementation
entire distribution range (Ortega et al. 2012), lead- of a marine zoning (Castrej
on and Charles 2013).
ing to a full fishery closure between 1995 and Co-M was institutionalized through two nested
2008. decision-making bodies: the Participatory Manage-
We analysed yellow clam information for the peri- ment Board (PMB) and the Institutional Manage-
ods 1982–1987 (before Co-M) and 1990–1994 ment Authority (IMA). Thus, local stakeholders
(after Co-M). Fishing metrics included clam abun- (i.e. fishers, touristic operators, naturalist guides,
dance and individual size, catch, fishing effort, CPUE and conservationists) were allowed to participate
and unit price paid to fishers at the beach. Abun- in the decision-making process along with the
dance and individual size data were obtained from Galapagos National Park Service (GNP) and the
seasonal surveys, following the methodology Ministry of the Environment, the institutions
described by Defeo (1996). Daily records of catch responsible for managing the GMR.
and effort per fisher and unit price were collected Sea cucumber abundance and individual size
through logbooks and interviews. A continuous spa- data were collected through population surveys
tial gradient in clam abundance precluded the possi- conducted by the GNP, Charles Darwin Founda-
bility of setting spatial controls (see also Castilla and tion and local users from 1999 to 2013, following
Defeo 2001). Therefore, Co-M performance was the methodology described by Toral-Granda and
assessed by testing hypotheses through time, using Mart
ınez (2004) and Wolff et al. (2012). Fishery-
temporal controls as predicting units. related information for the sea cucumber (1999–
2011) and spiny lobster (1997–2012) fisheries
was compiled from the Participatory Programme
Galapagos Islands (Ecuador): the sea cucumber
of Fisheries Monitoring and Research and from
Isostichopus fuscus and spiny lobsters Panulirus
Ram
ırez et al. (2012). Data were collected by
penicillatus and P. gracilis
interviewers and fishery observers at the three
The most important artisanal shellfisheries har- main ports of Galapagos (Puerto Ayora, Baquerizo
vested in Galapagos are the sea cucumber I. fuscus Moreno, and Villamil) on a daily basis along each
and spiny lobsters P. penicillatus and P. gracilis. fishing season. Fishing metrics included individual
These shellfish are harvested by hookah and skin- size (only for spiny lobster), catch, fishing effort,

180 © 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192
 Co-management in small-scale fisheries O Defeo et al.

CPUE, unit price paid to fishers at home port and and San Felipe, referred here as weakly managed
revenues per unit of effort. lobster fishing grounds, see details in Defeo and
Castilla 2005 and references therein) for four
fishing seasons comprised between 1998–99 and
Mexico: the spiny lobster Panulirus argus in
2001–02. A control–impact analysis was per-
Yucatan Peninsula
formed to assess Co-M performance by comparing
Panulirus argus is targeted by one of the most Punta Allen (co-managed site) with Chiquil
a and
important artisanal lobster fisheries in the world. San Felipe cooperatives (control sites).
In Mexico, state-authorized artisanal fishing coop-
eratives have had primacy to harvest spiny lob-
Analysis
sters and other high-valued coastal shellfishes
since the late 1940s (Table 1). One of these coop- Changes in fishery indicators associated with Co-M
eratives is ‘Vig
ıa Chico’, located at Punta Allen in implementation (abundance, individual size, land-
the Yucatan Peninsula, Mexico. It was formed by ings, fishing effort, CPUE, unit price and economic
49 fishers in 1969 to extract spiny lobsters at revenues per unit of effort) were estimated as
Ascension Bay, an area located inside the Sian response ratios of the mean and the variance
Ka’an Biosphere Reserve. Fishing operations are (Hedges et al. 1999). The response ratio of the
carried out exclusively by cooperative members mean gauged how the mean levels of the indica-
within a fishing ground of 850 km² granted by tors responded to Co-M implementation. The
the government. Spiny lobsters are harvested by response ratio of the variance measured changes
skin-divers, who use a hand net (locally known as in the interannual variability that coincided with
‘jamo’) to catch and keep lobster alive. The area Co-M implementation.
granted to Vig
ıa Chico cooperative has been We tested whether shellfishery stocks were
divided by members into individual ‘campos’ (fish- improved after Co-M implementation (before–after
ing grounds) that vary from 0.5 to 3.0 km2 (Seijo analysis) or when compared with open access sites
1993; Sosa-Cordero et al. 2008). The number of (control–impact analysis). Different strategies were
‘campos’ has varied through time between 150 set in each case, according to the nature, amount
(Seijo 1993) and 101 (Orensanz et al. 2013). Each and extent of the information gathered (see details
cooperative member has exclusive rights to deploy above for each fishery). ANOVAs were also employed
‘Cuban casitas’ (artificial shelters) and extract lob- to test the null hypothesis of absence of differences
ster within the assigned boundaries. Ownership or in indicators of Co-M success through time and/or
access rights to the most productive areas are space. ANCOVAs were used to test for differences in
given to those fishers with longer and continuous the unit price – catch relationship before and after
activity. Fishing grounds are inheritable and trans- Co-M implementation. The unit price was used as
ferable according to community-established rules, the dependent variable, landings as the covariate
including monetary payments and/or barter trans- and the fishing regime (before–after Co-M) as the
actions (Seijo 1993). Management regulations fixed factor. A Bartlett test was performed prior to
established by the Mexican government (e.g. mini- all analyses to test the assumption of homogeneity
mum legal size, fishing season) have been of variances among treatments. When data were
respected and enforced by cooperative members. heteroscedastic, transformations were carried out.
Furthermore, Vig
ıa Chico’s fishers have agreed
upon the implementation of additional well-defined
Results
internal rules to avoid illegal fishing (e.g. internal
extra penalties). Our long-term bioeconomic analysis of selected
Daily catch and effort data (ca. 43 000 records) Latin American shellfisheries showed that most
for the Punta Allen spiny lobster fishery were indicators were highly responsive to Co-M imple-
obtained from logbooks for the period 1986–2003. mentation (Fig. 1). After Co-M implementation (i)
CPUE (catch per trip), price paid to the fishers at abundance of yellow clam increased 108%,
home port and daily total revenues per fisher and whereas the sea cucumber decreased 70%; (ii)
fishing trip were also recorded. Catch and effort CPUE of loco, yellow clam and Galapagos spiny
information was also gathered for two nearby lobster increased 12, 66 and 9%, respectively,
cooperatives at the Yucatan Peninsula (Chiquil
a whereas the sea cucumber decreased 59%; (iii)

© 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192 181
Co-management in small-scale fisheries O Defeo et al.

landings and fishing effort showed in all cases a (Table 3) and eight of these significantly differed
drastic reduction (46–82% and 33–84%, respec- between fishery regimes (ANOVA: Table 2), whereas
tively); (iv) unit prices and revenues per unit of nine combinations indicated a 25% or greater
effort substantially increased (37–210% and 25– reduction in the mean following Co-M implemen-
805%, respectively); and (v) mean individual sizes tation (eight differed significantly between fishery
remained fairly constant. Before–after comparisons regimes: Table 2). Particularly notable was a large
through ANOVA roughly followed these patterns and consistent decrease in landings and fishing
(Table 2). effort following Co-M implementation, together
Overall, our results showed that 10 of 28 fish- with a marked increase in unit price and revenues
ery/indicator combinations had a 25% or greater per unit of effort (Tables 3 and S1). The response
change in the response ratios of the mean ratio of the variance showed large positive

Figure 1 Long-term mean (SE) response ratios (logarithmic basis) for seven bioeconomic indicators of Co-M success,
estimated for four Latin American shellfisheries. Log (response ratios) >0 ( ), that is untransformed values >1, implies
an increase in mean values of the indicators after the implementation of Co-M, whereas values <0 ( ) imply a decrease
in the mean. Number of years before and after Co-M implementation is given in Table 1.

Table 2 ANOVA results (F-ratios) for seven fishery indicators used to test the performance of Co-M in four small-scale
Latin American shellfisheries. Arrows indicate declining or increasing values in the indicators after Co-M
implementation, whereas = indicates no significant differences between fishery regimes (P > 0.05). *: P < 0.05;
**: P < 0.01; ***: P < 0.001. In the case of the Chilean loco fishery, only 1 year was available for the pre-Co-M (open
access) fishery phase at El Quisco (local level), and therefore, ANOVAs were only conducted for indicators at the national
level.

Metric

Fishery Abundance Size Landings Effort CPUE Unit price Revenues UE

Loco (Chile, country level) 72.65*** 48.69***

Yellow clam Uruguay 5.30* 0.06= 9.24** 247.23*** 19.83** 42.98*** 35.73**
Sea cucumber Galapagos 23.10** 4.67= 5.80* 2.95= 10.11** 12.21** 6.80*
Spiny lobster Galapagos 5.19= 16.03* 8.33* 0.18= 8.06* 1.09=

182 © 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192
 Co-management in small-scale fisheries O Defeo et al.

Table 3 Frequency of substantial (magnitude >25%) shifts in the mean or variance for seven fishery indicators used to
test the performance of Co-M in four small-scale Latin American shellfisheries. Figures in parentheses indicate the
numbers of each indicator that were statistically significant (P < 0.05), based on one-way ANOVAs. No: number of cases
used for assessment.

Mean Variance

Metric No. 25% increase 25% decrease 25% increase 25% decrease

Abundance 2 1 (1) 1 (1) 1 1

Size 3 0 (0) 0 (0) 3 0
Landings 4 0 (0) 4 (4) 0 4
Effort 3 0 (0) 3 (2) 1 2
CPUE 3 1 (1) 1 (1) 1 1
Unit Price 4 4 (4) 0 (0) 3 1
Revenues UE 4 4 (2) 0 (0) 2 0

changes (variance >25%) in 11 of the 28 fishery/ ative at Punta Allen (a co-managed lobster fish-
indicator combinations (notably in individual size, ery) were significantly higher than those estimated
unit price and revenues per unit of effort), whereas for the control sites Chiquil

a and San Felipe coop-
nine analyses showed a reduced variance in the eratives (2-way ANOVA: F2,62 = 64.51; P < 0.001,
co-managed phase (Tables 3 and S2). However, a Table S3). The ‘site x year’ interaction factor was
consistent response in the reduction of the vari- not significant, meaning that the effect of the fac-
ance was found only for landings. tor ‘site’ was constant through time, reflected in
Figure 2 shows CPUE variations through time consistently higher CPUE estimates at Punta Allen
as an indicator of Co-M success. CPUE increased (co-managed site) throughout the study period
after Co-M implementation in the loco (El Quisco, (Fig. 2e).
Chile: Fig. 2a) and yellow clam (Fig. 2b) fisheries, Relationships between catch and unit price
reaching values two times higher than in pre-Co- (Fig. 3) showed similar decreasing trends, but with
M years. In both fisheries, this trend was accompa- different interpretation among cases. In Chile (both
nied by a substantial decrease in fishing effort and at the national and cove-specific scales: Fig. 3a)
landings, which reached similar levels to the and Uruguay (Fig. 3b), prices increased at low
development phase of these fisheries, and a catch levels during the Co-M period. In these
marked increase in unit price (Fig. 1, Table S1). cases, low catches were accompanied with high
The Galapagos spiny lobster also showed a signifi- abundance (Fig. 1) and CPUE (Fig. 2). In Chile,
cant decrease in fishing effort and landings after loco’s catches were lowest after Co-M implementa-
Co-M implementation, as well as a significant tion, concurrently with the highest unit export
increase in unit price (Table 2). Even though prices (Fig. 3a). The Co-M phase (1994–2009),
annual average CPUE did not show significant dif- with a steeper slope, had higher prices than those
ferences between fishery regimes (Table 2), a sub- under open access (1981–1988), mainly at low
stantial and systematic increase of this indicator landing levels, denoting a very dissimilar expecta-
has been observed between 2006 and 2012 tion of unit prices for a given catch (ANCOVA:
(Fig. 2c). A very different scenario was found in F1,21 = 6.77; P < 0.05) and a willingness to pay
the sea cucumber fishery, where CPUE decreased higher prices during the Co-M phase. Indeed, dur-
43% after Co-M implementation (Fig. 2d), concur- ing the Co-M phase, the unit price (17 818 US
rently with a significant decrease in abundance $ t
1) was almost three times higher than before
and an increase in unit price (see ANOVAs in Co-M implementation (6292 US$ t
1). The long-
Table 2 and Figs 3d and 4a). term analysis at the scale of one Co-M area, El
Concerning spiny lobster fishery in the Yucatan Quisco (Central Chile: 33°230 S), mimics major
Peninsula (Mexico), our analysis showed that market macroscale trends: the price paid per loco
CPUE estimates obtained from Vig
ıa Chico cooper- at the Caleta was inversely related to the number

© 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192 183
Co-management in small-scale fisheries O Defeo et al.

of locos caught (Fig. 3a, right-hand-side panel). catches reached the lowest levels, whereas the
The yellow clam fishery in Uruguay showed a sim- unit price paid to fishers at the beach significantly
ilar pattern (Fig. 3b): after Co-M implementation, increased from 0.91
US$ kg
1 (before Co-M) to 1.70 US$ kg
1 (after
(a)
Co-M). The slope was steeper for the Co-M phase
than in the pre-Co-M period (ANCOVA: F1,5 = 16.90;
P < 0.01).
Following the same pattern found for Chile and
Uruguay, the unit price–catch relationship in the
spiny lobster Galapagos fishery showed a higher
price paid for a given catch level during the
Co-M phase (ANCOVA: F1,12 = 4.96; P < 0.05,
Fig. 3c). The same trend was observed in the sea
(b) cucumber Galapagos fishery (Fig. 3d). In this
case, unit prices paid for a sea cucumber
increased more than an order of magnitude
between 2002 (0.33 US$ ind
1) and 2011 (4.00
US$ ind
1). However, unlike Chile and Uruguay,
the exponential rise in unit price occurred con-
currently with a significant decrease in catch,
abundance and CPUE through time (Fig. 4a),
suggesting over-exploitation. This perception is
supported by five fishery closures implemented
(c)
since 2006 (Fig. 4a). Concerning the Galapagos
spiny lobster, the last period of the Co-M phase
showed a systematic increase in CPUE, together
with a stabilization of fishing effort and an
increase in unit prices (Fig. 4b). Taken together,
these trends determined the highest revenues per
unit effort since 1997.
Intra-annual patterns in the price–catch rela-
tionship for the Vig
ıa Chico cooperative at Punta
Allen (Mexico) also showed a monotonic decreas-
(d) ing trend: the average price paid per tonnes of
spiny lobster increased from the beginning (July)
to the end (February) of the fishing season, sug-
gesting short-term changes in price according to

Figure 2 Long-term variations in CPUE for five Latin

American shellfisheries: a) loco in El Quisco, Chile
(ind diver
1 h
1: mean  standard error); b) yellow clam
(e) in Uruguay (kg h
1 fisher
1: mean  standard error); c)
spiny lobster in Galapagos (kg tail diver
1 day
1); d) sea
cucumber in Galapagos (ind diver
1 h
1); and e) spiny
lobster in Mexico (kg boat
1 day
1). In the first four
cases, open access ( ), implementation ( ) and Co-M ( )
fishery phases are highlighted. In e), lobster CPUE
estimates obtained from daily records during four fishing
seasons in Punta Allen (Co-M site ), Chiquil
a and San
Felipe (control sites ) are shown. In the case of sea
cucumber, the fishery has been closed five times (2006,
2009, 2010, 2012 and 2013).

184 © 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192
 Co-management in small-scale fisheries O Defeo et al.

the magnitude of landings (Fig. 3e) and resource and Defeo 2003) define the upper limit of the
availability (considering CPUE as an index of relationship given by maximum unit prices P for
abundance, see Fig. 5). The points of the upper varying levels of lobster catch C (P =
boundary (a constraint envelope pattern, Caddy 33.10C
0.12; r2 = 0.67; P < 0.01). Most of these
points represent fishing activities developed either
at the beginning or at the end of the fishing sea-
son (Fig. 3e).
(a) Co-M was implemented in Punta Allen since
1969 (Table 1). Therefore, an additional way to
demonstrate its success is by looking at the long-
term performance of available indicators. An
analysis of ca. 43 000 daily records for the period
1986–2003 showed a fairly recurrent intra-
annual pattern in P. argus catch (Fig. 5a), fishing
effort (Fig. 5b) and CPUE (Fig. 5c). In 15 (fishing
effort) or in all (landings and CPUE) of 17 analy-
(b) sed years, these fishery variables consistently
peaked at the beginning of the fishing season
(July), decreasing exponentially until dropping
down to their lowest level at the end of the fishing
season (February). Only seasons 1994/1995 and
1995/1996 were exceptions to this pattern. No
significant differences in annual landings and fish-
ing effort were detected (ANOVAs, P > 0.05), reflect-
ing a remarkable long-term stability of these
(c)
indicators. CPUE showed significant differences
between years (ANOVA: F16,112 = 4.93, P < 0.001),
and multiple comparisons (Tukey HSD test)
revealed that only 13 of 136 paired tests showed
significant differences, with only three fishing sea-
sons (95/96, 97/98 and 02/03) having signifi-
cantly higher CPUE values than the remaining
ones (Fig. 5c).

(d)

Figure 3 Scatter diagrams for the relation between

catch and unit price in five Latin American shellfisheries:
a) loco, both at the country level (catch in t 103 and
price in US$ 103 t
1) and for the Co-M area El Quisco in
Central Chile (panel at the right-hand side: # of locos
103 and US$ loco
1); b) yellow clam (Uruguay),
expressed in tonnes and US$ kg
1; c-d) Galapagos spiny
lobster (t tails and US$ kg tail
1) and sea cucumber
(e) (catch in millions of individuals and price in US$ ind
1);
and e) spiny lobster (Punta Allen, Mexico). In a-d, open
access ( ), Co-M transition phase ( ) and Co-M ( )
fishery phases are highlighted. In e), the exponential
decreasing function for the spiny lobster in Mexico was
fitted only for empty circles ( ), which define the upper
limit of the ‘envelope’ between catch (t) and unit price
(US$ kg
1), representing maximum unit prices for
varying catch levels. Values within this ‘envelope’ ( )
represent suboptimal conditions (see text for details).

© 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192 185
Co-management in small-scale fisheries O Defeo et al.

(a)

(b)

Figure 4 Long-term variations in fishery indicators in Galapagos shellfisheries: a) sea cucumber and b) spiny lobster.
Closed seasons for the sea cucumber fishery are highlighted. All variables were standardized (z-scores) for clarity.

urchins (Loxechinus albus, Echinidae) and three

Discussion
species of keyhole limpets (Fissurella spp., Fissurel-
Our long-term analysis of multiple Co-M regimes lidae) were larger in Co-M areas (MEABRs) than
supports the hypothesis that Co-M can be an effec- in open access grounds (Castilla and Fern
andez
tive governance mode to sustain shellfisheries, 1998; Castilla and Defeo 2001; Gelcich et al.
leading to improved social–ecological fishery con- 2010). This provides well-grounded evidence that
ditions and greater production capacity provided the implementation of Co-M in combination with
that some key attributes are present. We described spatial property rights led to stability in shellfish
a common improvement of the main response individual sizes well above minimum legal sizes
variables relating to bioeconomic performance of and thus could be considered as an additional
these fisheries, specifically, increased population indicator of success.
abundance, higher CPUE, and higher unit prices The reduced variance in the response ratio of
and economic revenues at low landing and fishing landings denotes a reduction in the interannual
effort levels (Table S1). During the Co-M phase, variability of this indicator following Co-M imple-
the loco and yellow clam fisheries registered low mentation, which has key implications for fisheries
and stable catch levels, resembling initial phases of sustainability (Essington 2010) and market stabil-
fishery development. This fact, together with more ity and supply. This trend was observed jointly
temporally stable landings, suggests that successful with the stabilization of TACs and the compliance
Co-M have made these shellfisheries more predict- of fishers to catch quotas. Castilla et al. (1998)
able through time (Defeo and Castilla 2005; showed that Chilean shellfishers in open access
Essington 2010). These systematic indicator sites were unable to achieve their assigned quotas,
responses were the direct result of a set of man- whereas organized fishers in Co-M areas (‘caletas’)
agement measures agreed between local communi- not only completed the assigned quota, but also
ties and management authorities to sustain obtained significantly higher selling prices thanks
landing levels over time. Individual sizes did not to the reorganization of their harvesting and mar-
improve significantly following Co-M implementa- keting strategies (Defeo and Castilla 2005). Simi-
tion. However, short-term increases in individual larly, during the Co-M phase of the yellow clam
sizes were observed in Chilean shellfisheries during fishery in Uruguay, fishers achieved their assigned
Co-M implementation. For example, locos, sea individual quotas and the TAC set for the fishing

186 © 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192
 Co-management in small-scale fisheries O Defeo et al.

(a)

(b)

(c)

Figure 5 Monthly variations, July (J) to February (F), of (a) catch, (b) effort and (c) CPUE for the spiny lobster
(P. argus) fishery at Punta Allen (Mexico) for the fishing seasons between 1986/1987 and 2002/2003. Information is
based on daily records gathered from the Vig
ıa Chico cooperative.

season, avoiding quota overages (Castilla and Defeo By contrast, Co-M implementation was unable
2001). Viable explanations for this response include to prevent the collapse of the Galapagos sea
the end of the race to fish at Co-M sites, likely cucumber fishery. Several factors are responsible
because Co-M provided appropriate incentives lead- for this failure (Castrej
on 2011; Castrej
on and
ing to a strong sense of ownership and accountabil- Charles 2013), including the following: (i) weak
ity (Castilla 2010; Gelcich et al. 2010). The loco leadership, lack of social cohesion and poor orga-
and yellow clam fisheries provided a ‘response with nization of local fishing sector; (ii) lack of long-
experience’ (sensu Folke et al. 2003) to the over- term strategic planning and mechanisms for
exploitation crisis that occurred before Co-M precautionary and adaptive management; (iii)
implementation; that is, a social and institutional incapacity of the rights-based management system
learning has occurred based on previous experi- (i.e. licenses and fishing permits) to mitigate over-
ences, crises and social–ecological memory (Barthel exploitive fishing practices; and (iv) poor imple-
et al. 2010; Gelcich et al. 2013). In both fisheries, mentation, enforcement and compliance with
institutional learning was denoted by the imple- management regulations. The latter was encour-
mentation of management measures focused on aged by an anthropogenic Allee effect (sensu
stopping access to over-exploited fisheries, promot- Courchamp et al. 2006); that is, as resource abun-
ing efficient reorganization of harvesting and dance becomes scarcer, the willingness of external
marketing strategies, as well as the appropriate allo- agents, particularly Asian middlemen, to pay higher
cation of resources to enforce management interven- prices for sea cucumbers increased exponentially
tions, and to ensure effective coordination and (Fig. 4a). In this context, the Galapagos Co-M
collaboration with local actors (Gelcich et al. 2010). regime was unable to break down the pervasive

© 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192 187
Co-management in small-scale fisheries O Defeo et al.

partnership created between ‘roving bandits’ (sensu 4 years (2009–12), which were mainly reflected
Berkes et al. 2006) and local fishers since 1992, in the stabilization of fishing effort and the highest
which led to the intensification of illegal fishing and CPUE and revenues per unit effort values observed
to the collapse of the fishery in 2006 (Castrej
on since 1998, that is, since the beginning of the Co-
et al. 2014). This case-study illustrates how the M implementation phase. The moratorium on new
institutionalized inclusion of fishers as partners in entrants after 2002 also contributed to this recov-
the governance process (at least on paper) does not ery by slowing down the exponential growth in
necessarily produce positive fishery outcomes; that fishing effort. Nowadays, the only ones with the
is, Co-M is not a blueprint approach that can be right to reclaim their access to Galapagos fisheries
applied to all small-scale shellfisheries to enhance are the children of fishers. However, as most of
their governability. them are not interested in becoming fishers (Cas-
Most factors described above for the sea cucum- trej

on 2011), the number of license holders has
ber also affected negatively the Galapagos spiny been kept relatively stable. These results showed
lobster fishery from the late 1990s to the middle that the development of a Co-M approach in Gala-
2000s (i.e. during and after Co-M implementa- pagos shellfisheries is a work in progress and still
tion), leading to its over-exploitation (Fig. 2 and requires improvements and careful monitoring of
4b, and Castrej
on et al. 2014). This period coin- the observed long-term trends.
cided with the closure of the sea cucumber fishery The ‘Vig
ıa Chico’ Cooperative lobster fishery at
in 2006 (Fig. 4a). The declining profitability of Punta Allen has been the most productive cooper-
both fisheries resulted in a voluntary abandon- ative along the Yucatan Peninsula (Mexico) dur-
ment of fishing by ca. 294 fishers between 2005 ing the last decades and constitutes a well-
and 2008 (Castrej
on 2011; Ram
ırez et al. 2012). institutionalized and resilient Co-M system that
In the lobster fishery, this led to a 56% reduction emerged from the local community some 40 years
in fishing effort. This trend was intensified in ago (Castilla and Defeo 2001). This governance
2009 during the climax of the global financial cri- mode served to sustain a resilient small-scale shell-
sis, which contracted the consumption of lobsters fishery over time. Despite the metapopulation
in the United States and European Union (Monne- characteristics of the spiny lobster, the co-man-
reau and Helmsing 2011), resulting in a 32% drop aged fishery at Punta Allen showed a high persis-
in local prices. Consequently, total fishing effort tence and stability, reflected by the recurrent
and landings declined 20 and 23%, respectively, patterns in landings, fishing effort and CPUE esti-
between 2008 and 2009. Further, an El Ni~ no mated from 17 years of daily data. Individual lob-
event occurred simultaneously in 2009 (Defeo ster sizes harvested by the local fishery are larger
et al. 2013), having a positive impact on spiny than those harvested and commercialized by other
lobster 2 years later (see Figs. 2c and 3c). For cooperatives in the Yucatan Peninsula (Sosa-Cor-
example, CPUE and landings increased 91% and dero et al. 2008). Self-organization and a strong
102%, respectively, whereas fishing effort only social cohesion have been key drivers for the suc-
increased 6% between 2009 and 2011. A similar cess of this Co-M system, where coastal-parcelled
pattern was also observed 2 years after the 1997/ areas (TURFs) have been effectively allocated dur-
98 El Ni~ no (see Fig. 3c). In this sense, Defeo et al. ing the last five decades to families as inherited
(2013) documented a significant relationship owners for lobster extraction. The geographic iso-
between 2-years lagged production (linearly detr- lation also promoted a high motivation on a self-
ended catch series) and sea surface temperature help approach to community development, with
that explained 36% of the annual production reg- minimum government interventions (Defeo and
istered for the spiny lobster fishery. Therefore, the Castilla 2005). The strong collective organization,
remarkable and consistent CPUE recovery observed with penalties imposed by well-defined operational
between 2006 and 2012 (i.e. during the Co-M rules enforced and controlled by local fishers on
regime) could be attributed to the substantial this well-defined and isolated territorial permit,
reduction in fishing effort, together with the com- explains the long-term success of this fishery.
bined effect of market forces and favourable envi- The Punta Allen spiny lobster fishery was more
ronmental conditions (Defeo et al. 2013). This resilient than other analogous fishery systems in
combination of factors determined very stimulat- the Yucatan Peninsula, absorbing unexpected
ing trends in fishery indicators during the last shocks such as the Hurricane Gilbert (year 1988)

188 © 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192
 Co-management in small-scale fisheries O Defeo et al.

and the global financial crisis (Castrej

on and Defeo requires strong social leadership (Guti
errez et al.
2014), without changing in fundamental ways. 2011) and social systems that foster cooperation
The co-governance system enhanced its own and self-enforcement of social norms (Ostrom
capacity to cope with uncertainty and surprise by 2009).
mobilizing diverse sources of resilience that
retained essential structures, processes and feed-
Conclusions
backs (Castilla and Defeo 2001; Castrej
on and
Defeo 2014). For example, the devastation caused The shellfisheries of Chile, Uruguay and Mexico
by the Hurricane Gilbert encouraged fishers to evaluated here suggest that Co-M led to several
take preventive actions to strengthen the fishery positive effects, including the following: (i) stabil-
governance and the financial administration of ization of landings at levels similar to early phases
their cooperative (Carr 2007; UNDP 2012). Vig
ıa of exploitation; (ii) enhancement of bioeconomic
Chico’s fishers responded collectively by fine-tun- indicators, including abundance, individual size,
ing their fishing effort (e.g. daily fishing trips) CPUE and economic revenues per unit of effort;
according to resource abundance, and applying and (iii) reduced interannual variability in several
rigorous internal penalties to members that indicators, particularly in landings. Perceptions
infringed federal and internal management regula- and environmental awareness of fishers engaged
tions (Sosa-Cordero et al. 2008). Concerning the with the co-managed policy have changed, with
financial crisis, adaptive and successful community fishers themselves becoming stewards of their
responses included product and market diversifica- resources and ecosystems (Chapin et al. 2010;
tion, and the acquisition of the infrastructure, Gelcich et al. 2010, 2013). The joint implementa-
technology and expertise needed to increase the tion of catch shares (TURFs and community quo-
production and exportation of live lobsters to Asia tas) under a collaborative/voluntary community
and some European countries where lobster framework resulted in increasing profits and
demand was higher. This fishery was recently cer- resource abundance (Defeo and Castilla 2012),
tified by the Marine Stewardship Council (MSC) while providing additional biodiversity conserva-
ecolabelling programme, being one of the few tion benefits (Gelcich et al. 2012). Social attri-
Latin American artisanal fisheries that received butes, including leadership and social cohesion,
this international recognition. also contributed to success, as highlighted for
Fisheries Co-M has a better probability of suc- Chile (Gelcich et al. 2010), Uruguay and Mexico
cess when redundancy in management regulations (Castilla and Defeo 2001) and in Co-M systems
is in place (Caddy and Defeo 2003; Stefansson and throughout the world (Guti
errez et al. 2011).
Rosenberg 2005; Guti
errez et al. 2011). Our Despite the long-term success of co-managed
results demonstrate that Co-M is successful when shellfisheries, their social–ecological resilience and
combined with clear incentives, such as individual sustainability should not be taken for granted, as
or community quotas, TACs and spatial property demonstrated for the Galapagos sea cucumber fish-
rights (Chile, Uruguay and Mexico). Strategically ery. Co-M is not a panacea, that is, a blueprint for
sited MPAs could be an effective complement to a single governance mode that is applied to all
spatial property rights-based fisheries, increasing environmental problems (Ostrom et al. 2007). Co-
both fishery profits and abundance (Costello and M may work nicely under stable and predictable
Kaffine 2009). Co-managed TURFs also comple- conditions, but its resilience will be tested when
ment biodiversity objectives of MPAs. For example, subject to shocks, natural disasters and other types
Co-M areas in Chile registered higher species rich- of crises (Folke et al. 2003). Price shocks or rap-
ness and greater abundance of target benthic spe- idly emerging markets performing like ‘roving ban-
cies (e.g. loco, sea urchins, limpets and algae) and dits’ (Berkes et al. 2006) for sea cucumbers can
non-target sympatric fauna (including reef fish cascade from local to global environments (Ander-
species) than open access areas, leading to more son et al. 2010), and therefore, this problem is
diverse community assemblages (Gelcich et al. beyond the jurisdiction of national institutions.
2012). In addition, fishers have been able to diver- Future studies need to be undertaken for small-
sify their individual portfolios by harvesting a wide scale shellfisheries highly embedded in global mar-
variety of shellfishes within their Co-M areas, kets, such as in the loco and sea cucumber fisher-
using different fishing methods. Finally, Co-M ies, where price fluctuations strongly influence

© 2014 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 17, 176–192 189
Co-management in small-scale fisheries O Defeo et al.

local communities and their shellfisheries. This is Beck, M.W., Brumbaugh, R.D., Airoldi, L. et al. (2011) Oys-
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sustainability of social-ecological systems. Science 325, Table S1. Changes in mean estimates (%) of
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Ostrom, E., Janssen, M.A. and Anderies, J.M. (2007)
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(kg boat
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