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Benefits of No-take Zones for Belize and the Wider Caribbean Region


Abstract and Figures

Marine Protected Areas (MPAs) are now widely used for marine conservation and fisheries management, but the effects and benefits of MPAs may depend on what uses they allow or restrict. For many conservation and fisheries management goals, no-take zones or fully protected areas can provide the greatest benefits. As part of Belize’s expansion of its network of no-take zones (also referred to as replenishment zones) to include 10% of its marine territory, a global review of the benefits and potential social or economic costs of no-take zones was conducted. This review compiles and builds on over 20 years of research into the efficacy of no-take zones with respect to conservation, fishery, ecosystem function, social and economic benefits. The ecological and socio-economic factors that affect the ability of no-take zones to produce these benefits are also discussed. As part of this review, we examine the benefits provided by no-take zones in Belize and around the region. Our findings highlight numerous benefits from Belize and throughout the wider Caribbean region, including examples of how no-take zones may be effective for conservation and fishery management of key species like queen conch, Caribbean spiny lobster, and Nassau grouper. There is also evidence for the role of no-take zones in support of protecting or enhancing ecosystem function and coral reef resilience for Belize and across the region. While not all no-take zones will provide the same benefits and their effectiveness will be determined by various aspects of their design and implementation, well planned no-take zones should prove to be a valuable component of broader marine resource management for Belize and throughout the region.
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Proceedings of the 67th Gulf and Caribbean Fisheries Institute November 3 - 7, 2014 Christ Church, Barbados
Benefits of No-take Zones for Belize and the Wider Caribbean Region
Beneficios de Zonas sin Explotación de Belice y la Región del Gran Caribe
Avantages de ne pas Exploiter les Zones de Belize et de la Région des Caraïbes
1247 High Country Rd. Fayston, Vermont 05673 USA. *
2Wildlife Conservation Society, 1755 Coney Drive, Belize City, Belize.
Marine Protected Areas (MPAs) are now widely used for marine conservation and fisheries management, but the effects and
benefits of MPAs may depend on what uses they allow or restrict. For many conservation and fisheries management goals, no-take
zones or fully protected areas can provide the greatest benefits. As part of Belize’s expansion of its network of no-take zones (also
referred to as replenishment zones) to include 10% of its marine territory, a global review of the benefits and potential social or
economic costs of no-take zones was conducted. This review compiles and builds on over 20 years of research into the efficacy of
no-take zones with respect to conservation, fishery, ecosystem function, social and economic benefits. The ecological and socio-
economic factors that affect the ability of no-take zones to produce these benefits are also discussed. As part of this review, we
examine the benefits provided by no-take zones in Belize and around the region. Our findings highlight numerous benefits from
Belize and throughout the wider Caribbean region, including examples of how no-take zones may be effective for conservation and
fishery management of key species like queen conch, Caribbean spiny lobster, and Nassau grouper. There is also evidence for t he
role of no-take zones in support of protecting or enhancing ecosystem function and coral reef resilience for Belize and across t he
region. While not all no-take zones will provide the same benefits and their effectiveness will be determined by various aspects of
their design and implementation, well planned no-take zones should prove to be a valuable component of broader marine resource
management for Belize and throughout the region.
KEY WORDS: No-take zones; Marine Protected Areas, Belize
Throughout the Caribbean, coral reefs have been in decline, with widespread losses in coral cover due to localized
stresses and regional bleaching events and disease outbreaks (Jackson et al. 2014). Furthermore, loss of the sea urchin,
Diadema antillarum, and reduction of large bodied parrotfish species allows macroalgae to take over open space on reefs,
inhibiting coral growth and recruitment and further compromising the ability of coral reefs to recover from stresses ( Jackson
et al. 2001, Knowlton 2004, Mumby 2009). As a result, benthic communities of Caribbean reefs have changed dramatically
and structural engineers such as corals have been greatly reduced, leading to changes in habitat structure for fish and mobile
invertebrates that live on reefs (Alvarez-Filip et al. 2009). Declines are not limited to coral reefs, with significant losses in
other interconnected tropical coastal systems including seagrass and mangroves (Jackson et al. 2001, Alongi 2002, Orth et
al. 2006).
In response to these threats, marine resource managers are increasingly turning to ecosystem-based approaches to
management, including the use of marine protected areas (MPAs). MPAs are areas of the sea where some or all uses of
marine resources are restricted to enhance the management or conservation of them. MPAs may be created for a variety of
purposes – fisheries management, biodiversity conservation, recreational uses, or to address the conservation needs of
sensitive species or habitats (Sobel and Dahlgren 2004). As such, they may be created and administered by different natural
resource management authorities and under separate pieces of legislation in the same country, and the designation and use
of MPAs also differs between countries (e.g., Sobel and Dahlgren 2004, Dahlgren 2014).
Many MPAs are divided into zones with different levels of protection for resources. MPAs, or zones within MPAs that
have been shown to have the greatest positive effect on resources, are ones in which resource extraction, such as fishing, is
prohibited. These areas go by many different names but are commonly referred to in the scientific literature as no-take
zones or fully protected reserves. In Belize, the term replenishment zone is being adopted to refer to these areas to empha-
size the potential benefits that they provide rather than restrictions on uses of the areas.
In the wider Caribbean region, Belize has been a leader in the implementation of MPAs, where they have been created
by several different management authorities under separate pieces of legislation (Gibson et al. 2004). They may be designat-
ed as Marine Reserves under the responsibility of the Fisheries Department, or National Parks, Sanctuaries, or Monuments
under the responsibility of the Forest Department (Table 1). In addition, there are Marine and Forest Reserves under the
authority of both the Fisheries and Forest Departments. Many MPAs have co-management arrangements with local non-
governmental organizations. Within each of these areas, various uses are restricted, and zoning is often used to restrict use s
further in specific areas. Zones may vary among MPAs, but typically include General Use Zones, limited-use zones (e.g.
Conservation Zones) and no-take zones (i.e., some Conservation Zones and Preservation Zones; Table 2).
Belize is currently in the process of expanding its national network of no-take zones from a total of approximately 3%
of its marine territory to 10%. As part of this expansion effort a comprehensive review of the global literature related to t he
effects and benefits of no-take zones was conducted (Dahlgren 2014). While this review of these benefits is comprehensive
in its scope, it is not meant to be a complete review of all no-take zone literature. Here, we summarize findings from this
report using examples from Belize, with additional support from studies elsewhere in the wider Caribbean region. Benefits
are grouped into the following general categories:
Dahlgren, C. and A. Tewfik GCFI:67 (2015) Page 265
i) Conservation,
ii) Fisheries,
iii) Ecosystems and their function, and
iv) Socio-economic.
Because no-take zones prohibit extraction or consump-
tion of marine resources, one of their greatest documented
effects has been to increase the abundance, individual size,
and ultimately fecundity, of species targeted in fisheries.
The conservation of target species is critical in providing
other benefits, including benefits to fisheries, ecosystem
protection, and social and economic benefits. Thus, any
discussion of the benefits of no-take zones must begin with
the conservation of targeted species. Over the past 10 - 15
years, this effect has become well-established in the
literature and the subject of several review, synthesis and
meta-analysis studies (Halpern 2003, Micheli et al. 2004,
Lester et al. 2009, Molloy et al. 2009, Babcock et al. 2010).
This benefit well documented for queen conch (Lobatus
gigas), Caribbean spiny lobster (Panulirus argus) as well
as several species of grouper and snapper in Belize and
throughout the region.
The queen conch, Lobatus gigas (formerly Strombus
gigas), has shown positive responses to protection within
no-take zones throughout the wider Caribbean region
(Stoner and Ray 1996, Stoner et al. 1998, Béné and Tewfik
2003). In Belize’s MPA system, conch densities have been
monitored as part of the Long-Term Atoll Monitoring
Program (LAMP) within a no-take zone in Glover’s Reef
Marine Reserve and in areas open to fishing showed an
increase in adult conch density from 1998 through 2004 in
the no-take zone, which was not observed in fished areas
(Figure 1, Acosta 2006).
There is also a national conch measuring program
which is conducted in several marine reserves around the
country. Data from these surveys and LAMP surveys
provide a mixed success of no-take zones for protecting
conch populations, with several MPAs showing strong
positive effects of no-take zones on conch densities (Table
3). In at least one case, overfishing and poaching, particu-
larly from foreign fishers, were cited as reasons for the
failure for no-take zones to produce positive effects on
conch stocks (Chicas and Williams 2012).
Reports of greater conch densities in no-take zones are
of particular importance for the viability of the species due
to Allee effects in its reproduction. Recent work in The
Bahamas indicates that there was greater number of conch
in no-take zones and a 90% probability of successful
mating when densities were above 100 adult conch per
hectare in a no-take zone, while outside the no-take zone,
densities of 350 - 570 conch per hectare were needed for
reproduction to occur (Stoner et al. 2012a, Stoner et al.
2012b). This difference is believed to be due to selective
removal of larger individuals (i.e. older/thick lipped/more
mature) in fished areas leaving largely sexually immature
individuals. A related study found that current fisheries
regulations for conch focusing on a minimum shell length
(e.g., Belize) or simply the presence of a flared lip of the
shell (e.g., Bahamas) are inadequate to ensure that
individuals reach maturity before being harvested (Stoner
et al. 2012c). Shell length at maturity may vary, leading to
the capture of many juveniles that mature at larger sizes, or
the development of the flared shell lip occurs prior to
actual sexual maturity, with 50% of the population
reaching maturity only after the shell lip thickens to 24 - 26
mm, with a recommendation of a minimum lip thickness of
Table 1. List of MPAs of Belize with their management authority, total area and no-take Area
Protected Area Name Management/
Authority IUCN Category Marine area
(Km2) No-take area
Bacalar Chico National Park and Marine
Reserve Fisheries Dept. IV 52.8 9.2
Blue Hole Natural Monument Forest Dept. / BAS III 4.1 4.1
Caye Caulker Marine Reserve Fisheries Dept. /
FAMRACC VI 38.9 14
Caye Glory Spawning Aggregation Fisheries Dept. IV 5.5 5.5
Corozal Bay Forest Dept. IV 703.7 0
Gladden Spit & Silk Cayes Marine Reserve Fisheries Department/SEA IV 110.4 1.5
Glovers Reef Marine Reserve Fisheries Department IV 350.7 73.4
Half Moon Caye Natural Monument Forest Dept./BAS II 39.2 39.2
Hol Chan Marine Reserve Fisheries Dept. II 54.4 4.2
Laughing Bird Caye National Park Forest Department II 41 41
Nicholas Caye Spawning Aggregation Fisheries Dept. IV 6.7 6.7
Northern Glover's Spawning Aggregation Fisheries Dept. IV 6.2 6.2
Port Honduras Marine Reserve Fisheries Dept./TIDE IV 395 13.2
Rise and Fall Bank Spawning Aggregation Fisheries Dept. IV 17.2 17.2
Rocky Point Spawning Aggregation Fisheries Dept. IV 5.7 5.7
Sandbore Spawning Aggregation Fisheries Dept. IV 4.5 4.5
Sapodilla Cayes Marine Reserve Fisheries Dept./SEA IV 156.2 4.9
Seal Caye Spawning Aggregation Fisheries Dept. IV 6.5 6.5
South Pt. Lighthouse Spawning Aggregation Fisheries Dept. IV 5.3 5.3
South Water Caye Marine Reserve Fisheries Dept. IV 476.7 89.9
Swallow Caye Wildlife Sanctuary Forest Dept./FOSC IV 33.5 36.3
Turneffe Atoll Fisheries Dept. IV 1176.2 152.15
Totals 3690.4 540.7
Page 266 67th Gulf and Caribbean Fisheries Institute
14 mm for the fishery in the study area (Stoner et al.
2012c). Based on these findings, the vast majority of the
reproduction that supports Belize’s conch population may
be from no-take zones.
The LAMP surveys being implemented in MPAs in
Belize also record data on Caribbean spiny lobster
encounter frequency and sizes. These data show different
responses from MPAs, but in five of the seven MPAs for
which data is available, there were greater relative
abundances, frequency of occurrence and/or sizes reported
from no-take zones were greater than fished areas (Table
4). Elsewhere in the region studies of Caribbean spiny
lobster in no-take zones have shown greater densities
(Lipcius et al. 1997, Cox and Hunt 2005) particularly for
females (Bertelsen et al. 2004), and larger sizes than in
fished areas (Bertelsen and Cox 2001). Size may be
particularly important for lobster reproduction. For
example, males are typically larger than females under
natural conditions, but in fished populations, males and
females tend to be similar in size. When large males are not
present, females may not mate (Bertelsen and Matthews
2001, Butler et al. 2015). Furthermore, the size of the
spermatophore and amount of sperm cells deposited on the
female are correlated with the size of the male that deposits
them and has a strong influence on fertilization success
rates (MacDiarmid and Butler IV 1999, Butler et al. 2015).
In a large no-take zone in the Florida Keys, the sperm-to-
egg ratio was 40% higher than that found on average in a
fished population (MacDiarmid and Butler IV 1999, Butler
et al. 2011).
Similar to conch and lobster, no-take zones have been
shown to have significant positive impacts on commercial
finfish species of importance to Belize, including grouper
and snapper. Some of the earliest examples of the conser-
vation benefits of no-take zones for fish in the Caribbean
are from Belize with nearly half of all targeted species
observed in visual censuses of no-take areas of the Hol
Chan Marine Reserve having had a greater abundance,
size, or biomass than in ecologically similar fished areas,
with snappers showing the greatest difference in biomass
(Polunin and Roberts 1993). In fact, snapper biomass
increased by up to 220% in the Hol Chan Marine Reserve
within two years of its creation (Roberts 1995). Elsewhere,
Nassau grouper and other commercial species have greater
densities or biomass in no-take areas (Sluka et al. 1997,
Dahlgren 1999, Mumby et al. 2006). In Belize, Nassau
grouper spawning aggregations have decreased dramatical-
ly over the last 35 years as a result of fishing (Sala et al.
Table 2. Activities explicitly allowed (√) and prohibited (X) in various zones of Belize’s MPAs. (Hol Chan excluded -See text
for details.) Blank boxes indicate the activity is not addressed in relevant legislation. MR = Marine Reserve; CZ = Conser-
vation Zone.
Conservation Zone Activities General Fishing Subsistence
fishing by
Glover’s Reef MR CZ X
South Water Caye MR CZ X X X
Turneffe MR CZ I X
Turneffe MR CZ IIB *
Port Honduras MR CZ I X X X
Port Honduras MR CZ II X X
Bacalar Chico MR CZ I X X X
Bacalar Chico MR CZ II X **
Gladden Spit & Silk Cayes MR CZ I X X X
Gladden Spit & Silk Cayes MR CZ II*** X X
Caye Caulker MR CZ X X X
*no conch fishing; **only for catch-and-release tours; ***special regulations for whale shark viewing
Activities in Wilderness/Preservation Zones On/In Water activity (fishing,
diving, sportfishing, etc.) Motorized boats Any Vessels
Glover’s Reef MR X X
South Water Caye MR X X
Port Honduras MR X X
Bacalar Chico MR X X
Caye Caulker MR X X
Turneffe MR**** X X
**** New Turneffe MR includes spawning sites listed below
Spawning Aggregation Sites Any Fishing Some Traditional Fishing
Rocky Pt., Ambergis Caye X
Dog Flea Caye, Turneffe **** X
Caye Bokel, Turneffe**** X
Sandbore, Lighthouse Reef X
South Pt., Lighthouse Reef X
Emily X
Northern, Glover’s X
Gladden Spit
Rise and Fall Bank, Sapodilla Cayes
Nicholas Caye, Sapodilla Cayes
Seal Caye, Sapodilla Cayes
Dahlgren, C. and A. Tewfik GCFI:67 (2015) Page 267
2001). Inclusion of these spawning sites within MPAs and
designating them as seasonal fishing closures or full-time
no-take zones have improved the status of this species in
For fish that change sex like ecologically important
parrotfish and some grouper species, sex ratios and
population structure can differ between populations in no-
take zones and fished areas. For grouper that aggregate to
spawn, fishing can greatly reduce the number of males in
the population (Coleman et al. 1996). Furthermore, the
implementation of no-take zones can increase parrotfish
densities, the proportion of terminal phase males in the
population and the size at which females switched to
become terminal-phase males (Hawkins 2004). The effect
of no-take zones on population structure, however, may
take extended periods of time to develop (Molloy et al.
Fisheries Benefits
The increase in abundance and size of target species in
no-take zones can lead to improved fisheries in surround-
ing areas through larval replenishment or the “recruitment
effect” and the movement of juveniles and adults from the
no-take zone to fished areas or the “spillover effect”.
While both effects have been documented from various no-
take zones around the world (reviewed in Dahlgren 2014),
these effects have only been examined in a few studies in
the Caribbean region. For example, studies of conch and
lobster larval dispersal from a no-take area in The Bahamas
provide some evidence for potential recruitment effects
(Stoner et al. 1998, Lipcius et al. 2001), but no empirical
studies in the Caribbean to date have documented the
extent to which the recruitment effect enhances fisheries.
The spillover effect has been demonstrated in several
places by showing a decrease in density or landings of
fishery species with increasing distance from a no-take
zone (Halpern et al. 2010). For example, in Barbados, trap
catches of fish decreased with increasing distance from a
no-take zone (Chapman and Kramer 1999). Tagging
studies, however, suggest that many Caribbean reef fish
species have small home range sizes (e.g., Chapman and
Kramer 2000, Bolden 2001) and may rarely move across
no-take zone boundaries (Rakitin and Kramer 1996). For
example, some species that migrate to form spawning
aggregations like Nassau grouper have small home ranges
and may only leave no-take areas during spawning
migrations (e.g., Bolden 2000, Bolden 2001, Dahlgren et
al. submitted). Large parrotfish tagged in a small no-take
zone in Jamaica were captured more frequently outside the
no-take zone than smaller ones, suggesting spillover may
be the result of increased sizes in no-take zones (Munro
2000). The occurrence and extent of spillover may be
affected by habitat and seascape characteristics, such as
whether the no-take zone encompasses all of a habitat type,
whether the boundaries of zone cross continuous habitat
(Eristhee and Oxenford 2001), or if barriers to movement
Figure 1. Conch population size and structure within a no-
take zone and open fishing area in Glover’s Reef Marine
Reserve. Blank spaces represent missing surveys.
(Reprinted from Acosta 2006).
Table 3. Benefits of Belize’s no-take zones for queen conch, Lobatus gigas.
MPA Effect on conch Source
Gladden Spit and Silk Cayes Marine
Densities 2-4 times greater in no-take zones than unprotected or general use
zones Hagan (2010)
Laughing Bird Caye National Park Densities 2-4 times greater in no-take zones than unprotected or general use
zones Hagan (2010)
Bacalar Chico Marine Reserve Density in Preservation Zone greater than all other areas (2010-2012)
Density in Preservation zone more than tripled from 2010-2012 (Brown 2012)
Caye Caulker Marine Reserve 88% of conch observed were in no-take zones despite area only being 33%
of sampling (Cansino 2012)
Hol Chan Marine Reserve Density increased from 1998-2010, but then decreased Forman (2012)
Glover’s Reef Marine Reserve Most conch juveniles; changes over time nit assessed due to use of different
survey sites Chicas and Wil-
liams (2012)
Port Honduras Marine Reserve Conch populations increase from 2009-2010, particularly in no-take zone, but
then decrease in 2011. Poaching suspected Foley (2013)
Page 268 67th Gulf and Caribbean Fisheries Institute
exist (Tewfik and Béné 2003). Whether the effects of
spillover actually enhance fisheries depends upon whether
the increase in density, biomass, and ultimately catches of
target species around a no-take zone compensate for or
exceed the loss of abundance, biomass or potential
landings that would have been available to fishers if the no-
take zone had not been in place, and whether landings are
Ecosystem Benefits
Within no-take zones, the conservation of target
species and associated community structure, elimination of
bycatch, and prevention of habitat damage by fishing can
provide what may broadly be called ecosystem benefits.
Such benefits include the conservation of biodiversity and
support of ecosystem function. Ecosystem function
includes indirect effects that the no-take zone may have on
populations of non-target species to make the structure of
these populations and communities closer to pristine
conditions. Preserving ecosystem function also includes
enhancing or restoring ecosystem resistance or resilience to
various natural and anthropogenic threats and stressors,
including climate change.
The greater abundance of target species in no-take
zones can provide ecosystem benefits in two ways. First,
their occurrence in no-take zones and scarcity outside of
them can lead to direct increases of biodiversity and
redundancy within functional groups inside no-take zones
(Micheli et al. 2014). Their increased abundance within no-
take zones can also have indirect effects on other organ-
isms through competitive and trophic interactions. Within
no-take zones, the increase in target species can have
cascading effects throughout the ecosystem that may be
complex and interact (Kellner et al. 2010). In The Exuma
Cays Land and Sea Park, for example, increases in target
predator species may lead to decreases in their prey and
changes in benthic communities (Mumby et al. 2006,
Mumby et al. 2007, Harborne et al. 2009, Mumby et al.
2012). By maintaining high grazing rates, reducing coral
predators and maintaining cascading trophic interactions
that promote the survival of corals, no-take zones can
maintain natural ecosystem processes and facilitate reef
recovery from stresses. While no-take zones may do little
to prevent coral loss from global stressors like spikes in
water temperature associated with climate change, a global
analysis found that coral cover remained constant in MPAs
over time but declined in unprotected areas, suggesting
increased resilience of reefs within MPAs (Selig and Bruno
2010, Selig et al. 2012),
By eliminating fishing, no-take zones also reduce
bycatch, which can have a significant impact on non-target
species. Traps and nets all produce various levels of
bycatch mortality (Harper et al. 1994, Matthews and
Donahue 1996, Matthews et al. 2005). One study found the
abundance of non-target species on reefs, including
parrotfish and other ecologically important species, was
inversely related to fishing pressure across all sites
surveyed (Hawkins et al. 2007). In Curaçao, parrotfish
were the most abundant species caught in traditional fish
traps (Johnson 2010). While parrotfish are part of the
commercial catch in many parts of the Caribbean, it is now
illegal to fish them in Belize, although they figure as part
of the bycatch from trap fisheries. In Belize, parrotfish
biomass was significantly greater at sites within no-take
zones than in unprotected sites or General Use Zones of
MPAs (Dahlgren 2014). Similarly, in the Bahamas where
parrotfish are only just starting to be targeted, the biomass
of large parrotfish species was significantly greater in a no-
take area than unprotected reefs, with significant ecological
consequences to benthic communities (Mumby et al.
2006). Models of the impact of species taken as bycatch
indicate that their response to protection in no-take zones
may be dependent upon the interactions with those species
that are targeted (Reithe 2006, Kellner et al. 2010).
In addition to taking bycatch, many fishing gears cause
habitat damage. Studies of the use of fish and lobster traps
in the Caribbean region have found that they can signifi-
cantly reduce cover of live coral, damage coral, sponges
and gorgonians in reef habitats, and reduce shoot densities
in seagrass habitats (Sheridan et al. 2005, Uhrin et al. 2005,
Lewis et al. 2009). Habitat impacts may be greatest when
traps are moved by winds and waves during hurricanes and
even during winter cold front events (Lewis et al. 2009).
As traps deteriorate and break apart, the debris left behind
may account for up to one third of marine debris on reefs
and impact benthic communities (Chiappone et al. 2002,
Chiappone et al. 2005). Condos, casitas or shades, as they
are known in Belize, have an impact on benthic communi-
ties by reducing corals, gorgonians, and sponges for up to
seven meters away (Hunt 2011). Because shades often
concentrate lobsters and attract other species, they may
also affect the abundance of prey species (Eggleston et al.
1997, Nizinski 2007).
Table 4. Benefits of Belize’s no-take zones for Caribbean spiny lobster, Panulirus argus.
MPA Effect on lobster Source
Gladden Spit and Silk Cayes Marine
Reserve No effect on encounter rates or relative abundance Hagan (2010)
Laughing Bird Caye National Park Relative abundance increased over time.
Greater frequency of occurrence in no-take zones than other areas in 2008
and 2009
Hagan (2010)
Bacalar Chico Marine Reserve 2-12 times greater abundance in no-take zones than general use zones at
close of 2012 lobster season Brown (2012)
Caye Caulker Marine Reserve Too few lobster reported in any zone for analysis Cansino (2012)
Hol Chan Marine Reserve 4-5 times greater abundance in no-take zones than general use zones Forman (2012)
Port Honduras Marine Reserve No difference among zones until 2012 when more reported in no-take zones
than general use zone. No-take zone typically has larger individuals Foley (2013)
Glovers Reef Marine Reserve Average size nearly 2 times as large in no-take zone as general use zone Chicas and
Williams (2012)
Dahlgren, C. and A. Tewfik GCFI:67 (2015) Page 269
Social and Economic Benefits
The increase in target species populations, their
support of fisheries, and their role in maintaining ecosys-
tem function can all provide social and economic benefits
to people. In Belize, for example, the increase in conch
stocks within no-take zones has reached levels to meet the
overall national target set for fulfilling CITES non-
detriment finding requirements, enabling Belize’s conch
exports to continue without CITES’ proscription or other
intervention. If Belize’s conch management were deemed
by CITES (through the CITES Animals and Standing
Committees) not to meet the requirements of the treaty,
exports could be suspended by CITES, and this would limit
the fishery to the demands of the limited domestic market,
with obvious economic consequences (Acosta 2006, Smith
et al. 2008).
In addition to supporting fisheries and facilitating
management, no-take zones may also provide local
communities with non-extractive economic benefits
through tourism and other activities. This is particularly
important for a country like Belize, where nature-based
tourism has been shown to benefit local economies and
generate local support for conservation (e.g., Lindberg et
al. 1996). The value to tourism of no-take zones and other
MPAs derives from the improved quality and quantity of
fish and benthic communities (Wielgus et al. 2003). In
coral reef systems, the increase in abundance and size of
fish and also trophic cascades that improve coral health
often support high levels of dive tourism. Surveys of diving
within tropical MPAs around the world show that there is a
strong preference by divers for reefs with high coral cover
and diversity, as well as high fish abundances and diversity
(Rudd and Tupper 2002, Wielgus et al. 2010). An econom-
ic valuation of the Bonaire Marine Park, for example,
indicated that the park brought in US$4.8 million each year
in scuba diving revenues (Dixon et al. 2000).
The value of protecting species within no-take zones
may outweigh the value of those species in fisheries (Sala
et al. 2001). In Belize, fisheries revenues from inside
Glover’s Reef Marine Reserve were estimated at US$0.7 -
1.1 million for 2007, as compared with US$3.8 - 5.6
million from reef-related tourism (Cooper et al. 2009).
Furthermore, Belize’s coral reefs and mangroves contribute
US$14 - 16 million annually to fisheries and approximately
ten times that amount, US$150 - 196 million, to tourism
(Cooper et al. 2009). Both fishery and tourism value,
however, was far outweighed by the economic contribution
that these habitats provide in terms of shoreline protection,
with damage avoided due to these habitats valued at
US$231 - 347 million annually (Cooper et al. 2009). Thus,
maintaining the health of these ecosystems to preserve their
structure is critical. Improved function of ecosystems
within no-take zones suggest that they may be essential to
these ecosystem services.
Clearly not all no-take zones will provide the same
benefits. The performance of no-take zones will vary based
on the size, shape and spacing of the no-take zone (Halpern
and Warner 2003, Gaines et al. 2010), as well as the
relationship of various habitats in a seascape within no-take
zones and their surrounding area (Grober-Dunsmore et al.
2007, Nagelkerken et al. 2012) and the permeability of no-
take zone boundaries (Eristhee and Oxenford 2001, Tewfik
and Béné 2003, Bartholomew et al. 2007). The degree of
compliance with no-take regulations is also important to
the success of the no-take zone for provision of benefits
(e.g., Francini-Filho and de Moura 2008). The process by
which stakeholders are involved in the designation of no-
take zones (Suman et al. 1999), the uses allowed within
(Thurstan et al. 2012), and the management context of the
no-take zones, such as the Managed Access system being
implemented in Belize’s MPA network (Foley 2013, WCS
2013), also contribute to the success of no-take zones for
providing benefits. It is also important to recognize that
there may be short-term costs to fishers associated with
creating no-take zones due to reduced availability of
resources, but these short-term costs should be small
compared to longer-term economic benefits (Smith et al.
2010) and multitude of other benefits provided (Dahlgren
While not all no-take zones will provide the same
benefits, there is strong evidence from Belize and the wider
Caribbean that well designed no-take zones with a high
level of compliance can provide a wide range of benefits
for conservation, fisheries management, ecosystem
resilience, and people living in coastal communities.
Examples from around the world indicate that these
benefits can be even more extensive than those reported
from the Caribbean to date, and that further, long-term
studies of no-take zones in Belize and the wider Caribbean
may reveal additional benefits (Dahlgren 2014). For a more
extensive review of this topic with examples from around
the world, we refer readers to the full report on the benefits
of no-take zones that was conducted as part of the no-take
zone expansion effort in Belize (Dahlgren 2014).
This manuscript was based on a report developed for the Wildlife
Conservation Society, which is available at:
Data from Belize no-take zones was provided by the Belize Fisheries
Department, Healthy Reefs Initiative, Toledo Institute for Development
and Environment, and Southern Environmental Association. Preparation
of this report was facilitated by R. Lewis and valuable comments to an
earlier draft of the report were provided by A. Brautigam and J. Gibson.
Funding for the preparation of the manuscript was provided by the
Wildlife Conservation Society.
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... no-take areas). In Belize, the term "replenishment zone" has recently been adopted to replace "no-take zone" to emphasize the overall objective of such closed areas to enhance small-scale fisheries-based livelihoods both within the RZ and in surrounding areas that are fished (Dahlgren, 2014;Dahlgren and Tewfik, 2015;Tewfik et al., 2017). RZs include 13 fish spawning aggregation sites where almost all fishing is banned (Burns and Tewfik, 2016) and several national parks (e.g. ...
... Laughing Bird Caye) and wildlife sanctuaries (e.g. Swallow Caye) that have protected marine elements (Gibson et al., 2004;Dahlgren and Tewfik, 2015;Cox et al., 2017) ( Figure 2). Most recently, a nationwide network of TURFs (i.e. ...
... The GSSCMR has a focus on the protection of multi-species fish spawning aggregation sites and associated whale shark visits to feeding grounds (Gibson et al., 2004). No effect of management zones on encounter rates or relative abundance of spiny lobster has been observed at GSSCMR (Dahlgren and Tewfik, 2015) possibly owing to the small size of the RZ. (iii) Laughing Bird Caye National Park (LBCNP, established in 1991), which encompasses a single 41 km 2 RZ, did experience some relative lobster abundance increases over time (Dahlgren and Tewfik, 2015). ...
In Belize, the commercial harvest of spiny lobsters has occurred for approximately 100 years, provides critical livelihoods, and is the primary seafood export. We determined the first empirical estimate of size at maturity in Belize as well as eight fishery status indicators on several fishing grounds. The carapace lengths (CLs) at 50% maturity varied between males (98 mm) and females (86 mm) and are higher than the existing legal minimum of 76 mm. Time series analysis indicated decreasing proportions of mature individuals, decreasing size, and low spawning potential ratios (SPR) as well as high fishing mortality within fishing grounds. The pattern of decline in population status indicators across fishing grounds is consistent with a historical expansion of effort from north to south and offshore. Many indicators of population status within fishing grounds were improved with increasing area of replenishment zone and opposite to the historical expansion. However, overfishing is a problem across all areas examined. An increase in the legal minimum CL to 86 mm and examination of a maximum size limit will likely have significant positive effects on productivity and SPR, respectively, as well as protecting the pivotal role of spiny lobsters within the ecosystem.
... This concept has been adopted by marine resource management to identify and protect habitats that preserve species and provide sources of replenishment to fisheries beyond no-take areas (Rowley 1994, McClanahan and Mangi 2000, Roberts et al. 2001. In Belize, a broad network of marine reserves has been established to protect environmental health and support socio-economic needs and where the extraction of Queen conch (Lobatus gigas) by free-diving fishers provides significant revenue (Acosta 2006, Babcock et al. 2015, Dahlgren and Tewfik 2015. In order to understand the distribution of conch resources, especially mega-spawners (large, mature and highly fecund individuals) (Bertelsen and Matthews 2001, Stoner et al. 2012, Hixon et al. 2014, and support the overall management of the fishery, we surveyed a number of habitats across Glover's Reef Marine Reserve in May and June of 2015. ...
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Conference Paper
A natural refuge provides an inaccessible area that can protect populations from exploitation (Karpov et al. 1998, Tyler et al. 2009). This concept has been adopted by marine resource management to identify and protect habitats that preserve species and provide sources of replenishment to fisheries beyond no-take areas (Rowley 1994, McClanahan and Mangi 2000, Roberts et al. 2001). In Belize, a broad network of marine reserves has been established to protect environmental health and support socioeconomic needs and where the extraction of Queen conch (Lobatus gigas) by free-diving fishers provides significant revenue (Acosta 2006, Babcock et al. 2015, Dahlgren and Tewfik 2015). In order to understand the distribution of conch resources, especially mega-spawners (large, mature and highly fecund individuals) (Bertelsen and Matthews 2001, Stoner et al. 2012, Hixon et al. 2014), and support the overall management of the fishery, we surveyed a number of habitats across Glover's Reef Marine Reserve in May and June of 2015. In addition, we conducted assessments of shells and extracted soft tissues, with fishers, to determine the maturity of harvested conch with reference to three important indicators (Froese 2004). Fifty percent maturity was established at 5 mm shell lip thickness with a corresponding 160 g market clean meat mass (Babcock et al. 2015). Existing regulations consist of a minimum shell length of 178 mm (7 inches), no minimum shell lip thickness lip and a market clean meat mass of 85 g (3 ounces). Optimal size range and mega-spawner minimum, both at 100% maturity, were calculated to be 11 to 13 mm and 14 mm shell lip thickness respectively (Babcock et al. 2015). Although the densities of conch did vary between habitats and management zones (Figure 1), our assessments revealed that most harvested conch were juveniles or sub-adults (lip thickness < 5 mm) (Babcock et al. 2015). Immature conch were most abundant in shallow (1.6 +/ 0.4 m) patch reef (N = 11) and sand flat (n = 14) sites (Figure 2) where fishing effort is concentrated and matches the known life history for this species (Stoner and Sandt 1992). In contrast high proportions of mega-spawners occurred in deeper fore-reef (13.3 +/-0.6 m, n = 12) and seagrass (10.2 +/-1.2 m, n = 13) sites around patch reefs (Figure 2). These habitats, outside no-take zones, may provide additional refuges for mega-spawners given more limited accessibility by free-divers due to depth as well as high turbidity (seagrass) and wave exposure (fore-reef) (Karpov et al. 1998, Tewfik 2014). However, these refuges are subject to harvest and should be considered for the application of specific management measures given the low overall density of adults and their importance to the maintenance of recruitment. One such measure could be the total prohibition of fishing conch outside the reef crest thereby protecting fore-reef refuges where the largest individual mega-spawners were found.
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Technical Report
Abstract The increase in negative impacts due to unsustainable tourism practices, overfishing, marine pollution and global climate change has made coral reefs one of the world’s most threatened ecosystems. In order to improve the resilience of reefs to global changes, local stressors need to be managed. Historically, marine protected areas (MPAs) have been one of the most commonly applied tools for coral reef conservation, with variable success in restoring fisheries or benefitting reef health. Bacalar Chico Marine Reserve (BCMR) was established in northern Belize in 1996 as a small multiple-use MPA. In order to evaluate the management effectiveness of BCMR, dive surveys following the Mesoamerican Barrier Reef System Synoptic Monitoring Program (MBRS-SMP) protocol were conducted at ten sites within the reserve and two control sites outside the reserve over a five-year period (2011–15). Additional surveys were conducted on queen conch (Lobatus gigas) and Caribbean spiny lobster (Panulirus argus), both important commercial species in Belize, along with opportunistic sightings of megafauna recorded during routine activities. Results show that coral reef health in BCMR is ranked at Poor to Critical status under the Simplified Integrated Reef Health Index (SIRHI) scale. This ranking was based on several factors, including a significant decline in fish biomass between 2011–15, with Critical commercial fish biomass and herbivorous fish biomass in 2015. Although hard coral cover increased from Poor in 2011 to Fair in 2015 it was lower than the 2015 national average (15%), and fleshy macroalgae cover was higher than the 2015 national average (24%). The abundance of lobster and queen conch varied across management zones but with no significant difference between the years. Five species of ray, two species of shark and three species of marine turtle were sighted in BCMR between 2011–15; there was no significant difference per zone for elasmobranch sightings, and turtle sightings only differed significantly between reef types. There was minimal or no measurable influence of the different management zones on coral reef condition or the populations of commercial and endangered species. On the basis of these results, a reassessment of the management approach for BCMR is strongly recommended, including a review of the location of management zones, the implementation of an updated management plan, strengthening the enforcement of reserve regulations, and improving ridge-to-reef management focusing on land-based pollution within the Northern Belize Coastal Complex.
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In the Florida Keys, traps for spiny lobsters (also known as Caribbean spiny lobster) Panulirus argus are often deployed in seagrass beds. Given that several hundred thousand traps may be deployed in one fishing season, the possibility exists for significant impacts to seagrass resources. The question was whether standard fishing practices observed in the fishery actually resulted in injmies to seagrass. This study was designed to measure the degree of injury to seagrass as a function of trap deployment duration (soak time) and habitat type (seagrass species) and the recovery of seagrass following trap removal. Aspects of the deployment and retrieval process were not examined. Sampling grids composed of 30 3-m x 3-m squares were arbitrarily established within each of three monospecific seagrassbeds (two of Thalassia testudinum and one of Syringodium filiforme) near Marathon, Florida. Five squares within each grid remained trap-free (controls) while the remaining squares each received a single trap. Five traps from each grid were randomly removed at each of five soak times (ranging from 1 to 24 weeks). Immediately before deployment and following trap removal, seagrass short shoot densities were recorded and compared among controls and treatments. Both seagrass species exhibited significantly decreased shoot densities after 6-week and 24-week soak times. Thalassia testudinum densities within the 6-week and 24-week treatments had returned to control densities 4 months after trap removal, while densities of S. filiforme remained significantly decreased at the end of 24 weeks. We conclude that traps must be recovered within a 6-week period, beyond which injury to seagrass beds is predicted, with long lasting effects to beds of S. filifonne. Within the limits of these testing parameters, it appears that standard fishing practices (typically < 5-week soak time) should not result in a significant injury to seagrass beds in the Florida Keys.
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Populations of benthic species that produce pelagic larvae are sustained through a complex interaction of factors, including larval supply, variable transport mechanisms, and a host of postsettlement processes that affect differential recruitment and abundance. We report distributional data for the larvae, juveniles, adults, and a time-averaged index of fishery yield (shell middens) of the economically important marine gastropod Strombus gigas (queen conch) in the Exuma Sound, Bahamas. All life history stages and the fishery yields were heterogeneously distributed around this semienclosed system, with higher densities of benthic stages in the northern part of the sound than in the south and east. Distribution of shell middens closely reflected abundance patterns of shallow-water juvenile aggregations and abundance of adults in depth-stratified surveys; therefore, midden distribution provided a good indicator of long-term productivity around the periphery of the sound. Although patterns of fishery productivity around the system were closely related to both juvenile and adult distributions, and density of newly-hatched larvae reflected the distribution of adults and shell middens, as would be expected, benthic stages and the fishery yields were completely decoupled from the abundance of settlement-stage larvae. When transplants of newly settled conch were made to four seagrass sites in the eastern Exuma Sound with characteristics typical of conch nurseries, low growth rates resulted in all but one location. All of these results suggest that conch abundance and distribution in Exuma Sound is determined in the benthos, either during settlement or in the first year of postsettlement life. Therefore, although larval supply has been shown to influence benthic recruitment on a small scale (i.e., size and location of juvenile aggregations), and juvenile populations will always depend upon a reliable source of competent larvae, high quality habitat plays an equally important role in the recruitment of this important fishery resource.
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Effectiveness of a marine protected area (MPA) in supporting fisheries productivity depends upon replenishment patterns, both in supplying recruits to surrounding fished areas and having a sustainable spawning stock in the MPA. Surveys for queen conch Strombus gigas were made in 2011 at 2 locations in the Exuma Cays, The Bahamas, for direct comparison with surveys conducted during the early 1990s at Warderick Wells (WW) near the center of the Exuma Cays Land and Sea Park (ECLSP) and at a fished site near Lee Stocking Island (LSI). There was no change in adult conch density and abundance in the shallow bank environment at LSI where numbers were already low in 1991, but numbers declined 91% in the deeper shelf waters. At WW, the adult population declined 69% on the bank and 6% on the island shelf. Unlike observations made in the 1990s, queen conch reproductive behavior near LSI is now rare. Average age of adult conch (indicated by shell thickness) at LSI decreased significantly during the 20 yr period between surveys, while average age increased at WW and juvenile abundance decreased. These results show that the LSI population is being overfished and the WW population is senescing because of low recruitment. In 2011, the ECLSP continued to be an important source of larvae for downstream populations because of abundant spawners in the shelf environment. However, it is clear that the reserve is not self-sustaining for queen conch, and sustainable fishing in the Exuma Cays will depend upon a network of MPAs along with other management measures to reduce fishing mortality.
Lobster populations in southern Florida fall into three size-classes: less than 10 km2, 10-100 km2, and more than 100 km2. Databases spanning the past 30 years are being reexamined to investigate the relationship of size of sanctuary on size structure, density, and fecundity of populations of spiny lobster Panulirus argus (also known as Caribbean spiny lobster). The density of the lobster population in small sanctuaries (established in 1997) has not changed; however, a few larger males may be protected. In the medium-sized sanctuary, Western Sambo Ecological Reserve (WSER), the density of male lobsters has roughly doubled and that of females has quadrupled. When the large sanctuary, Dry Tortugas National Park (DTNP), was established in 1974, large females (>120 mm carapace length) did not bear eggs, perhaps because there were no large males there. Today, all large females bear eggs and contribute 31% of the total fecundity. Fecundity estimates for area and season are difficult to calculate because they are sensitive to size distribution, fecundity-to-size relationships, and other factors. Our preliminary estimate is that 28 million eggs are produced per season per ha in the fore reef of DTNP, 21 million in WSER, 18 million in small sanctuary preservation areas, and 14 million in the Florida Keys fishery.
The average size of spiny lobsters (Decapoda; Palinuridae) has decreased worldwide over the past few decades. Market forces coupled with minimum size limits compel fishers to target the largest individuals. Males are targeted disproportionately as a consequence of sexual dimorphism in spiny lobster size (i.e. males grow larger than females) and because of protections for ovigerous females. Therefore, overexploitation of males has led to sperm limitation in several decapod populations with serious repercussions for reproductive success. In the Caribbean spiny lobster, Panulirus argus, little is known about the effect of reduced male size on fertilization success or the role that individual size plays in gamete and larval quality. We conducted a series of laboratory experiments to test the relationship between male size and spermatophore production over multiple mating events and to determine whether spermatophore reduction and female size affected fertilization success or larval attributes in P. argus in the Florida Keys, FL (USA). We found that over consecutive matings, larger males consistently produced spermatophores of a greater weight and area than smaller males, although size-specific differences in sperm cell density were undetected and probably obscured by high variance in the data. Where spermatophores were experimentally reduced to mimic the decline in spermatophore size with declining male size, fertilization success (the number of fertilized eggs/total number of eggs extruded) declined, indicating that sperm availability is indeed limited. No maternal size effects on egg size or quality (C:N ratio) or larval quality (size, swimming speed, mortality) were observed. Our results demonstrate the importance of maintaining large males in populations of P. argus to ensure fertilization success and caution against their overexploitation through fishing, which may severely reduce reproductive success and thus population sustainability.
Widespread use of minimally selective fish traps has contributed to the overfishing of Caribbean coral reefs. Traps typically target high-value fish such as groupers (Serranidae and Epinephelidae) and snappers (Lutjanidae), but they also have high bycatch of ecologically important herbivores (parrotfish (Scaridae) and surgeonfish (Acanthuridae)) and non-target species. One strategy for reducing this bycatch is to retrofit traps with rectangular escape gaps that allow juveniles and narrow-bodied species to escape; yet the effectiveness of these gaps has not been thoroughly tested. On the shallow reefs of Curacao, Netherlands Antilles, I compared the catch of traditional Antillean chevron traps (the control) to the catch of traps with short escape gaps (20 x 2.5 cm), traps with tall escape gaps (40 x 2.5 cm), and traps with a panel of large aperture mesh. With data from 190 24-h trap sets, the mean number of fish caught was 11.84 in control traps, 4.88 in short gap traps, 4.43 in tall gap traps, and 0.34 in large mesh traps. Compared to controls, traps with short or tall gaps caught significantly fewer bycatch fish (-74 and -80% respectively), key herbivores (-58 and -50% respectively), and butterflyfish (Chaetodontidae; -90 and -98% respectively). The mean length of captured fish was significantly greater in gap traps because juveniles were able to escape via the gaps. Escape gaps reduce neither the catch of high-value fish, nor the total market value of the catch. Therefore, using escape gaps could make trap fishing more sustainable without reducing fishermen's revenues.
Marine fishery reserves (MFR's) have been set aside in coastal areas throughout the world with the hope of reversing population decreases commonly observed in many marine resources. In this study, a comparison of population structure of the commercially important gastropod Strombus gigas, queen conch, was made between a fished area and an MFR in the Exuma Cays, central Bahamas. There were 31 times more adult conch on the shallow (<5 m) Great Bahama Bank in the MFR, and in a survey at 7 depth intervals (to 30 m) on the island shelf in the Exuma Sound, mean adult density was always higher in the MFR, by as much as 15 times. Shell length and lip-thickness measurements indicated that adults in the MFR migrate with age from bank nursery sites into deeper sound water, whereas those on the bank in the fished area were harvested before reaching water sufficiently deep to protect them from free-diving fishermen. Although sparsely distributed juveniles in shallow-water (<15 m) habitats of the sound were the primary source of adults in the fished area, large juvenile aggregations on the bank also contributed to the deep-water adult stock in the MFR. Total larval densities in the MFR were frequently an order of magnitude higher than those found in the fished area, and densities of late-stage larvae were 4 to 17 times higher. Because the surface current along the Exuma Cays shelf flows to the northwest, late-stage larvae found inside the reserve must have been spawned outside the reserve; thus the high densities of juvenile and adult conch are the result of natural accumulation of larvae in the area, as well as the result of protection from fishing. Although the fate of larvae dispersed from the reserve is uncertain, it is likely that high numbers of reproductive stock and larvae in the reserve have a significant positive effect on populations in the northern Exuma Sound. Designs of reserves that consider ontogenetic requirements of the target species and strategic locations for larval production, import, export, and metapopulation dynamics will optimize fishery benefits for the many marine vertebrate and invertebrate species that possess pelagic larvae.