DRAFT – Please do not copy or cite without permission of the authors.
XIIIth Conference of the European Association of Fisheries Economists
18-20th April 2001, Salerno, Italy
Cost and benefits of implementing a marine reserve
facing prey-predator interactions1
Jean Boncoeur*, Frédérique Alban*, Olivier Guyader** and Olivier Thébaud**
* Université de Bretagne Occidentale, Centre de Droit et d’Economie de la Mer, Brest, France
(e-mail : Jean.Boncoeur@univ-brest.fr , Frederique.Alban@univ-brest.fr )
** IFREMER, Service d’Economie Maritime, Brest, France
(e-mail : Olivier.Guyader@ifremer.fr , Olivier.Thebaud@ifremer.fr)
Abstract
This paper deals with the economic effects of implementing a marine reserve when the impacts of the
reserve on both commercial fisheries and recreational activities – through changes in the ecosystem are taken into account. The issue is illustrated by the example of creating a no-take zone in part of a
region inhabited by two interacting stocks : a stock of preys (fish) and a stock of predators (seals).
While the former is targeted by commercial fishing, the latter is a potential basis for the development
of a commercial activity related to ecotourism. Analysis is conducted with the help of a bio-economic
model combining the features of marine reserve modelling and of multi-species modelling. Following
a description of the model, results of several simulations run using the software STELLA  are
presented. These show that creating a marine reserve may have more complex economic implications
than predicted in studies centred exclusively on commercial fisheries. The exact nature of the impacts
will depend on the ecological dynamics at play, the dynamics of commercial fisheries, and the
dynamics of the other activities, particularly recreational, that depend upon the ecosystem.
Keywords
Marine protected areas, multi-species interactions, ecotourism, bio-economic modelling.
1
The research presented in this paper is part of a research program concerning the economic aspects of
implementing a national park in the Iroise sea, funded by the French Ministry of Environment (MATE,
programme espaces protégés) and by the French National Program of Research on Coastal Environment (PNEC,
CNRS-IFREMER-IRD). It also takes advantage of the participation of the authors in a European Union funded
research program concerning the economic value of fishing exclusion zones (VALFEZ. project QLK5-CT199901271).
1
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Introduction
Various and sometimes contradictory achievements are expected from the creation of marine reserves
(Hoagland et al., 1995; Murray et al., 1999; Shackle et al., 1995). The objectives pursued can
generally be classified under one of the following three categories: ecosystem preservation, the
management of commercial fisheries, and/or the development of recreational activities.
At such a general level, the degree of compatibility between the three objectives is difficult to assess.
It is bound to vary from case to case, depending on the exact conditions under which the plan to create
a marine reserve is being discussed. In some cases, the pursuit of these objectives can lead to conflicts
between the various economic agents involved in the creation of a protected area (Dixon et al. 1993;
Polunin et al., 2000). This makes it important to develop tools which allow the overall assessment of
the consequences of creating a marine reserve, taking into account the distribution of impacts amongst
the various categories of activities and economic agents involved.
The fast-growing literature on marine reserves has largely centred on the issue of commercial
fisheries management (see e.g. Bohnsack, 1993, 1996; Gubbay, 1996; Roberts et al., 1993; Somerton
et al., 1984). Bohnsack (1990) gives a comprehensive list of the potential biological effects that may
be anticipated from the establishment of a marine reserve, which includes: (1) protection of spawning
biomass, (2) providing a recruitment source for surrounding areas, (3) supplemental restocking of
fished areas through emigration, (4) maintenance of natural population age structure, (5) maintenance
of areas of undisturbed habitat, and (6) insurance against management failures in fished areas.
These biological effects are expected to make fish stocks more resilient to exploitation, thereby
reducing the risk of stock collapse (Carr et al., 1993; Dugan et al., 1993; Guénette et al, 1998; Plan
Development Team, 1990; Rowley, 1994). Marine reserves may in addition be regarded as a means of
dealing with the uncertainties related to stock assessment and effort control in fisheries management
(Clark, 1996; Lauck et al. 1998).
A number of studies have focused on the development of biological models (see e.g. DeMartini, 1993;
Guénette et al., 1999; Polacheck, 1990; Sladek Nowlis et al., 1999) and bio-economic models of
marine reserves (Anderson, 2000; Arnason, 2000; Conrad, 1999; Hannesson, 1998, 2000; Pezzey et
al., 2000; Sanchirico et al., 2000; Sumaila, 1998)2. Parallel to this theoretical work, studies have dealt
with applying the theory to empirical case studies (Holland et al., 1996; Holland, 2000; Whitmarsh et
al., 2000). These studies show that the actual consequences of a reserve depend on factors such as its
size, the characteristics of the ecosystem where the reserve is established (in particular the mobility of
the fish stocks it shelters), the dynamics of fishing fleets, and the management policy in the
surrounding areas.
In comparison, economists have granted relatively less attention to the other categories of objectives
that can be pursued in the implementation of marine reserves. In particular, changes in ecosystem
structure and functioning following the effective implementation of a marine reserve are bound to
affect the recreational uses, both extractive and non-extractive, which can develop in these areas. This
must be included in an overall assessment of the economic consequences of creating a marine reserve.
While various studies have considered this issue, particularly in tropical regions (see e.g. Agardy,
1993; Buerger et al., 2000; Davis et al, 1995; Ditton et al., 1986; Kenchington, 1991), only a few deal
fully with the economic evaluation of recreational uses (see e.g. Dixon, 1993).
The purpose of this paper is to present a simple simulation model of the economic consequences of
implementing a marine reserve in a predator-prey system in which (i) the stock of predators (e.g.
2
In part, these studies take their inspiration from research in the ecology of metapopulation harvesting (see e.g.
Bulte et al., 1999; Roberts, 1998; Sanchirico et al., 1999, 2000b; Tuck et al., 1994) and in the spatial dynamics
of fishing fleets (see e.g. Allen et al., 1985; Eales et al., 1986; Hilborn et al., 1992; Wilson et al., 1999).
2
DRAFT – Please do not copy or cite without permission of the authors.
seals) is liable to increase due to the rise in food supplies provided by the fishing ban, while (ii) its
presence in the region is an important input in the development of local ecotourism (e.g. seal
watching)3. The model allows the simulation of the impacts of various policies (in terms of reserve
size) on the economic rents generated by both commercial fishing and the ecotourism industry.
The paper is organised as follows. First the structure of the model is presented, stressing its role as a
heuristic device allowing the exploration of some of the indirect consequences of creating a reserve
when several stocks and several human activities are interacting in the region under consideration.
Second, results of simulation runs carried out using the software STELLA are used to discuss the
implications of creating a marine reserve in such a system.
1. Description of the model
The model presented here combines previous work related to the modelling of marine reserves on the
one hand, and to the modelling of multi-species harvesting on the other. Assumptions as regards the
consequences for a commercial fishery of establishing a marine reserve are taken from Hannesson
(1998), while predator-prey interactions are modelled following Flaaten (1989)4. An additional
feature of the model is the inclusion of a demand function for ecotourism activities, which allows the
determination of economic rents derived from non-extractive use of the ecosystem.
The region under survey is assumed to be homogenous from an ecological point of view and to be
pertinent for the purpose of managing a fishery. It shelters two stocks: a stock of preys (e.g. fish) and
a stock of predators (e.g. seals), named respectively F and M. The region is split into two sub-regions
of varying sizes: a marine reserve and a zone open to fishing, named respectively 1 and 2. The
proportion of the total region closed to fishing is  with 0 <  < 1.
The dynamics of both stocks are modelled as follows5:
(1)
dX 1
X F1
 rF .X F1.(1 )  T   . X F 1. X M
dt
 . X F max
(2)
dX 2
X F2
 rF .X F2 .(1 )  T   . X F 2 . X M  YF 2
dt
(1   ). X F max
(3)
dX M
X M
 rM .X M .(1 )
dt
X F1  X F 2
3
. The choice of this example was motivated by the existence of a debate on a such an issue in the context of the
forthcoming creation of a marine national park in the Iroise sea (ICES VIIe), a coastal sea west of Brittany
(France). This area is characterised by a great variety of marine living resources (Hily et al., 1999) and shelters a
small colony of seals which, local fishermen fear, might develop as a consequence of a part or total fishing ban
within the limits of the park. On the other hand, this possible development is regarded as an opportunity of new
incomes in a region visited by many tourists (Anon., 1999).
4
Supriatna et al. (1999) develop a mathematical model for a two-patch predator-prey system as a generalisation
of single-species harvesting theory. Their approach is in terms of dynamic programming and Lagrange
multipliers.
5
Simple logistic models are used here. For a modified version of this model, see Pezzey et al. (2000). Guénette
and Pitcher (1999) use an aged structured model.
3
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With:
XFi
the fraction of the fish stock biomass in sub-region i, i = 1, 2 (where 1 is the marine
reserve and 2 is the fishing zone)
XM
the seal stock biomass
rF
the intrinsic growth rate of the fish stock biomass
rM
the intrinsic growth rate of the seal stock biomass
XFmax
the maximum fish stock biomass of the total region under study
T
the net transfer of biomass from the marine reserve to the fishing grounds

the instantaneous fish mortality rate due to predation by seals

the equilibrium ratio of fish-to-seal biomass
YF2
the catch of fish in the region open to fishing
Only the stock of fish is targeted by commercial fishing6, with the mortality rate of fish due to fishing
directly proportional to fishing effort. Fishing effort is assumed to be fixed (at varying levels), e.g.
through a system of licenses.
The mortality rate of fish due to predation by seals is assumed to be directly proportional to the
abundance of seals, while the area’s carrying capacity for seals is assumed to be directly proportional
to the abundance of fish.
Seals are assumed to be ubiquitous throughout the whole region. The net transfer of fish biomass from
the reserve to the fishing grounds, T, is assumed to be proportional to the relative size of fish biomass
to carrying capacity in each sub-region7:
(4)
T   ..(1   ).( DF1  DF 2 )   X F1   ( X F1  X F 2 )
with:
DF1
Density in the reserve
DF2
Density in the fishing zone

Mobility coefficient for the fish stock (0 <  < 1).
and DF 1 
X F1
;
X F max
DF 2 
XF2
1   . X F max
Catch is assumed proportional to the density of fish biomass in the sub-region open to fishing:
6
It is assumed here that the predator species is not subject to harvesting, hence that any economic value attached
to its presence in the area under study is derived exclusively from non-extractive uses. This is coherent with the
objective of the paper. Various alternatives could be considered: the predator species could also be harvested by
a commercial fishery; by a recreational fishery; or by both. These alternatives will not be considered here.
7
Since the area is considered homogeneous in ecological terms, this is equivalent to assuming that fish migration
takes place depending on relative biomass density between the two areas. Following Hannesson (1998), it is
equivalent to assuming that T = s (DF1 - DF2 ) with s = ..(1 -  ) which is the migration function used in many
studies of population dynamics in patchy environments (see e.g. Sanchirico et al., 1999; Wilson et al., 2000,
Tuck et al., 1994; Roberts, 1998; Hilborn et al., 1992 ).
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DRAFT – Please do not copy or cite without permission of the authors.
(5)
YF 
q.EF . X F 2
1   . A
with:
EF
A measure of nominal fishing effort
Q
A normalised catchability coefficient
A
The total area of the region under study
As noted by Arnason (2000), with a model of perfect diffusion of fish biomass between sub-regions,
fish density inside and outside the marine reserve will always be the same, and the exclusion of
fishing fleets from part of the overall region will have no impact on catchability. This would not be
the case if less than perfect diffusion was observed.
Ecotourism in the region is assumed to depend directly on the presence of the predator (seals), being
based on a commercial activity of seal watching. For a given price, the demand for ecotourism8 is
modelled as an non linear increasing function of the abundance of seals in the area and of the level of
effort devoted by the local industry in developing this activity EM:
(6)
YM  a. X M .EM
b
c
where a, b, c are strictly positive parameters.
The net benefits derived from the activities of commercial fishing and ecotourism are defined as
follows:
(7)
R j  Pj .Y j  C j .E j
with:
Pj
The average unit revenue devoted to activity j (j = F , M)
Cj
The unit cost of effort devoted to activity j. (j = F , M)
and the prices of fish and ecotourism services, and the costs of production factors are exogenous
parameters.
The system thus modelled achieves an equilibrium state for:
(8)
dX F 1 dX F 2 dX M


0
dt
dt
dt
2. Simulation results and discussion.
Simulation experiments with the model were carried out using the software STELLA. Because of
the heuristic character of this model, simulation results are only qualitative, and were obtained with
the following parameter and initial state variable values:
8
Other forms for the demand function for seal watching could be tested. The present form assumes an increasing
relation between the demand for ecotouristic services and the stock of seals. An alternative would be to use a
logistic form for the function, with demand remaining at a low level for very low levels of seal biomass, and
stabilising for high levels of seal biomass.
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State variables
Initial value
Parameters
Value
XF,0
XF1,0
XF2,0
XM,0
750
.XF,0
(1-).XF,0
XF,0 / 
rF
rM
0,3
0.05
0,2
0,0025
0,001
10
1
0,8
0,2
1
1000
40
6
4
4
1

q


a
b
c
A
XFmax
EM
PM
CM
PF
CF
Simulations were run for varying levels of fishing effort (between 0 and 300) and sizes of the marine
reserve (between 0 and 90% of the region), with or without the presence of seals and associated
ecotourism activity in the system9.
Simulations results presented in the following paragraphs concern steady-state values achieved for
fixed values of fishing effort and reserve size. First, results when no prey-predator relation is included
are presented. Second, the impacts of introducing seals and seal-watching activities in the system are
discussed.
2.1.
Reserve effects without prey-predator interaction
Figure 1 presents the evolution of equilibrium surplus production of fish biomass when there is no
predator in the region for different levels of fishing effort and varying reserve sizes. Sustainable catch
is in this case equal to this surplus production. If the regulator actually controls effort and aims at
maximising sustainable catches, the preferred option will be to restrict effort to low levels without any
reserve10. For higher levels of effort and where effort is difficult to control (e.g. because of a lack of
information, or for political and social reasons), implementing a reserve may become a second-best
option (Bohnsack, 1990; Lauck et al., 1998).
Increasing the size of the reserve will have two main effects: it will restrict the area available for
fishing, but it will increase surplus production available for catch due to fish migration effects. As
fishing effort grows, sustainable catches tend to rely exclusively on the net transfer of fish biomass
from the reserve. For high levels of fishing effort, the marine reserve becomes an essential source of
biomass for the fishery.
9
Simulation experiments were executed with a delta time of 0,25 and a number of time steps sufficient for the
system to achieve a stable state for each fishing effort/reserve size combination. The average number of time
steps needed for the system to stabilise was approximately 130.
10
Under the restrictive hypothesis of the deterministic model used here.
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DRAFT – Please do not copy or cite without permission of the authors.
80
70
60
reserve = 0%
50
reserve = 30%
40
30
reserve = 60%
reserve = 90%
20
10
30
0
25
0
20
0
15
0
50
10
0
0
0
Total surplus of biomass
Figure 1 - Total surplus production of fish biomass as a function of fishing effort, for varying
reserve sizes
Fishing effort
Figure 2 presents the same simulation results in terms of economic rents derived from the fishery. The
same effects can be observed. Again with a capacity to control effort, the regulator will not use the
reserve as a management tool if the objective is to extract maximum rents from the fishery. Where
controlling effort is difficult, the reserve may appear as an interesting second-best management tool11.
Implementing a reserve will have two main economic consequences. First, fishing will remain
profitable for higher levels of fishing effort with a reserve of intermediate sizes. Second, with a large
reserve, such economic benefits may be lost due to an important restriction on the catch area.
Figure 2 - Fishery rent as a function of fishing effort, for varying reserve sizes (without seals)
300
100
reserve = 0%
30
0
25
0
20
0
15
0
reserve = 60%
-100
10
0
reserve = 30%
50
0
0
fishing rent
200
reserve = 90%
-200
-300
-400
Fishing effort
2.2.
Consequences of the prey-predator interaction
When fishermen compete with seals for the resource, surplus production is absorbed in part by
fishermen as catch, and in part by seals as food. The impact of introducing a predator is thus to reduce
equilibrium levels of fish catch and producer surplus for a given level of effort.
11
The possibility that the marine reserve allows protection of critical habitats and/or prevents risks of stock
collapse is of course another argument in favour of no-take zones, which cannot be assessed using this model.
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Figure 3 presents the changes in surplus production of fish biomass as a function of fishing effort for
varying reserves sizes. Simulations were run with a positive initial seal stock, hence a surplus
production of fish is observed in the absence of fishing effort.
The figure illustrates the relative impacts of fishers and seals on the system. The larger the reserve,
the weaker the impact of fishing effort on total surplus production: while seals are ubiquitous,
fishermen are confined to the zone open to fishing.
Figure 3 - Total surplus of biomass as a function of fishing effort , according to the reserve size
(With seals)
Total surplus of biomass
80
70
60
reserve = 0%
50
reserve = 30%
40
reserve = 60%
30
reserve = 90%
20
10
30
0
25
0
20
0
15
0
10
0
50
0
0
Fishing effort
By restricting access of fishermen to fish stocks, larger marine reserves will favour predation of fish
by seals to the detriment of fishermen. Figure 4 shows how the presence of a predator can reduce the
advantages of the marine reserve as a tool to improve the economic situation of the fishery, where the
predator is itself of no commercial value to the fishery. The exact nature of the impact varies
according to the size of the marine reserve.
Figure 4 - Sharing of biomass surplus between fishers and seals for two sizes of the reserve
reserve size=30%
reserve size=90%
Fleet catch
Fleet Catch
Seal predation
Fishing effort
2.3.
300
275
250
225
200
175
150
125
75
100
50 75 100 125 150 175 200 225 250 275 300
50
25
0
0
70
60
50
40
30
20
10
0
25
80
70
60
50
40
30
20
10
0
Surplus of biomass
Surplus of biomass
Seal predation
Fishing effort
Taking ecotourism into account
Figure 6 below presents results of simulations taking ecotourism into account, with a fixed level of
fishing effort and varying reserve sizes. In this case, total economic rent is the sum of fishery rent and
the economic rent derived from ecotourism in the region.
Fishing effort is assumed to be fixed at its open access equilibrium level when all the region is open
to fishing, i.e. no economic rent is extracted from the fishery for a reserve size of 0. Economic
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parameters of the model have been chosen in such a way that ecotourism becomes a profitable activity
when at least 10% of the region is designated as a marine reserve.
Figure 5 - Economic rent as a function of the reserve size, for a constant level of fishing effort
(E=104)
300
250
Economic rent
200
150
Fishery rent without seals
Fishery rent with seals
Ecotourism rent
Total economic rent
100
50
0
-50 0
0,1
0,2
0,3
0,4
0,5
0,6
0,7
0,8
0,9
1
-100
-150
Reserve size
With these assumptions, the steady-state economic rent derived from the fishery increases with the
size of the reserve from 0 to a maximum beyond which the restriction of fishing area more than
compensates the biological effects of implementing a no-take zone. In order to illustrate the
implications of having a predator in the system, fishery rent as a function of reserve size for the same
level of fishing effort is also shown in the figure. The impact of seals is clearly to reduce the benefits
derived by fishermen from the marine reserve.
However, the increase in seal biomass associated to larger reserve sizes also entails greater potential
for the development of ecotourism in the region. By assumption, the difference between total
economic rent derived from the management region and fishery rent grows with the size of the area
designated as a reserve, following the increase in the associated economic rent derived from seal
watching.
The figure thus illustrates the trade-offs which may be encountered when more than one activity
dependent upon the region in which a marine reserve is created are considered. In the case discussed
here, the maximum increase in fishery rent derived from implementing a reserve, in comparison to
what would occur under open access equilibrium, is achieved for a reserve size of approximately
30%. The maximum total economic rent derived from implementing a reserve is slightly higher, and is
achieved for a reserve size of approximately 55%. In other words, designing a marine reserve (with a
fixed level of fishing effort) so as to maximise total economic benefits will entail an opportunity cost
for the fishery.
With large reserve sizes, the benefits from ecotourism will in fact more than compensate for the
opportunity costs supported by fishermen. This raises the issue of the distributional implications of
marine reserve policies, whose benefits and costs may not accrue to the same economic agents.
Conclusion
The aim of the simulation model presented in this paper is to develop a heuristic approach to the
economics of marine reserves, with a multi-species perspective and taking non-extractive uses of
marine ecosystems explicitly into account.
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Simulation results show the complexity of relations between the creation of a reserve and the various
economic costs and benefits derived from the ecosystem it shelters, in such a model. If creating a
marine reserve appears as potentially beneficial from the perspective of ecotourism, its impacts on the
fishery are more ambiguous. On one hand, creating a reserve restricts the fishing grounds, to the
benefit of predators in the system. On the other hand, for a range of reserve sizes and a range of
fishing effort levels, it allows the fishery to improve its economic situation in comparison to a
situation without a reserve, due to the biological consequences of stock protection.
The degree of improvement will depend on the biological parameters chosen in the model,
particularly fish mobility and growth, and rates of predation. Of course, it will only be possible if, for
technical and/or institutional reasons, fishing effort remains fixed at the same level once the marine
reserve is created, i.e. no additional fishing effort enters the fishery. Where this is the case,
maximising the total economic rent derived from the creation of a marine protected area may require a
reserve size which is different from the reserve size which would bring the largest improvement in the
situation of the fishery, hence a the need to balance the different sectoral objectives pursued by policy
makers.
The possibility that the predator species in this model be itself harvested for commercial purposes, or
that its non-use value be an important element in the overall economic value of the marine ecosystem,
have not been considered in this paper. Their inclusion in the model would surely lead to characterise
another set of trade-offs which are bound to be at the centre of marine reserve policies.
10
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References
AGARDY, M. T. (1993). “Accommodating Ecotourism in Multiple Use Planning of Coastal and
Marine Protected Areas.” Ocean and Coastal Management 20: 219-239.
ALLEN, P. M. et J. M. McGLADE (1986). “Dynamics of discovery and exploitation: the case of the
Scotian Shelf Groundfish fisheries.” Canadian Journal of Fisheries and Aquatic Sciences 43: 11871199.
ANDERSON, L. G. (2000). Marine Reserves: a closer look at what they can accomplish. 10th
Biennal Conference of IIFET, 10-14 July, Corvallis, Oregon, USA.
ANON. (1999) Etude économique des activités liées à la mer d’Iroise. ADEUPa Brest / Portances /
SAFI, Brest (3 volumes).
ARNASON, R. (2000). “Marine reserves: is there an economic justification ?” Conference on the
economic of marine protected areas at the UBC Fisheries Center, Vancouver, July 6-7.
BOHNSACK, J. A. (1990),The potential of marine fishery reserves for reef fish managment in the
U.S Southern Atlantic. NOAA Tech. Memo. NMFS-SEFC-261, 40pp.
BOHNSACK, J. A. (1993). “Marine reserves: They enhance fisheries, reduce conflicts and protect
resources.” Oceanus 36(3): 63-71.
BOHNSACK, J. A. (1996). “Marine reserves, zoning, and the future of fishery management.”
Fisheries 21(9): 14-16.
BONCOEUR, J., ALBAN, F., et al., 2000, Activités halieutiques et activités récréatives dans le cadre
d’un espace à protéger: le cas du Parc National de la Mer d’Iroise, rapport intermédiaire, Programme
National “Environnement Côtier”, CEDEM, IFREMER, Brest, 213pp.
BUERGER, R., J. HILL, et al. (2000). “The impact of recreation on Barrier Island: a case study on
Masonboro Island.” Coastal Management 28(3): 249-259.
BULTE, E. H. et G. CORNELIS van KOOTEN (1999). “Metapopulation dynamics and stochastic
bioeconomic modeling.” Ecological Economics 30(2): 293-299.
CARR, M. H. and D. C. REED (1993). “Conceptual issues relevant to marine harvest refuges:
examples from temperate reef fishes.” Canadian Journal of Fisheries and Aquatic Sciences 50: 20192028.
CLARK, C. W. (1996). “Marine reserves and the precautionary management of fisheries.” Ecological
Applications 6(2): 369-370.
CONRAD, J. M. (1999). “The bioeconomics of marine sanctuaries.” Journal of Bioeconomics 1: 205217.
DAVIS, D., V. HARRIOTT, et al. (1995). “Conflicts in a marine protected area: Scuba divers,
economics, ecology and management in Julian Rocks Aquatic Reserve.” Australian Parks and
Recreation 31(1): 29-35.
11
DRAFT – Please do not copy or cite without permission of the authors.
DeMARTINI, E. E. (1993). “Modelling the potential of fishery reserves for managing Pacific coral
reef fishes.” Fishery Bulletin 91: 414-427.
DITTON, R. B. et M. L. MILLER (1986). “Coastal recreation: an emerging area of managerial
concern and research opportunity.” Leisure Sciences 8(3): 223-239.
DIXON, J. A. (1993). “Economic benefits of marine protected areas.” Oceanus 36(3): 35-40.
DIXON, J. A., et al. (1993). “Meeting ecological and economic goals : marine park in the Caribbean.”
Ambio 22(2-3): 117-125.
DUGAN, J. E. and G. E. DAVIS (1993). “Application of marine refugia to coastal fisheries
management.” Canadian Journal of Fisheries and Aquatic Sciences 50: 2029-2042.
EALES, J. et J. E. WILEN (1986). “An examination of fishing location choice in the Pink Shrimp
fishery.” Marine Resource Economics 2(4): 331-351.
FLAATEN, O. (1989) « The economics of predator-prey harvesting » in P.A. Neher et al. (eds) Rights
Based Fishing, Kluwer Academic Publishers: 485-503.
GUBBAY, S. (1996). Marine Refuges: the next step for nature conservation and fisheries
management in the North-East Atlantic?, A report for WWF-UK: 22p.
GUÉNETTE S. and PITCHER T. (1999) « An age structured model showing the benefits of marine
reserves in controlling overexploitation ». Fisheries Research, 39: 295-303.
GUÉNETTE, S., LAUCK, T. and CLARK, C.W. (1998) « Marine reserves : from Beverton and Holt
to the present ». Reviews in Fish Biology and Fisheries, 8: 251-272.
HANNESSON, R. (1998). “Marine reserves: what would they accomplish?” Marine Resource
Economics 13: 159-170.
HANNESSON, R. (2000). “The economics of marine reserves”, Conference on the economic of
marine protected areas at the UBC Fisheries Center, Vancouver, July 6-7.
HILBORN, R. and R.-B. KENNEDY (1992). “Spatial pattern in catch rates : a test of economic
theory.” Bulletin of Mathematical Biology 54(2/3): 263-273.
HILY, C. et al. (1999) Environnement naturel de l’Iroise. Bilan des connaissances et intérêt
patrimonial. Université de Bretagne Occidentale / Ministère de l’Aménagement du Territoire et de
l’Environnement, Brest (2 volumes).
HOAGLAND, P., KAORU, Y. and BROADUS, J.M. (1995) A methodological review of net benefit
evaluation for marine reserves. The World Bank, Environment Department, paper n°027, 69p.
HOLLAND, D. S. (2000). “A bioeconomic model of marine sanctuaries on Georges Bank.” Canadian
Journal of Fisheries and Aquatic Sciences 57: 1307-1319.
HOLLAND, D. S. and R. J. BRAZEE (1996). “Marine reserves for fisheries management.” Marine
Resource Economics 11: 157-171.
KENCHINGTON, R. (1991). “Tourism development in the Great Barrier Reef Marine Park.” Ocean
and Shoreline Management 15: 57-78.
12
DRAFT – Please do not copy or cite without permission of the authors.
LAUCK, T., CLARK, C.W., MANGEL, M. and MUNRO, G.R. (1998) « Implementing the
precautionary principle in fisheries management through marine reserves ». Ecol. Appl. 8 (Suppl. 1),:
S72-S78.
MURRAY, S.N. et al. (1999) « No-take reserve networks : sustaining fishery populations and marine
ecosystems ». Fisheries Management, 24(11): 11-25.
PEZZEY, J., ROBERTS, C. and URDAL, B. (2000) « A simple bioeconomic model of a marine
reserve ». Ecological Economics, 33: 77-91.
Plan Development Team (1990). The potential of marine fishery reserves for reef fish management in
the U.S. Southern Atlantic. Miami, NOAA Technical Memorandum NMFS-SEFC-261: 40p.
POLACHECK, T. (1990). “Year around closed areas as a management tool.” Natural Resource
Modeling 4(3): 327-354.
POLUNIN, N. V. C. et al. (2000). Ecological and social impacts in planning Caribbean marine
reserves. Final Technical Report R6783, DFID, University of Newcastle: 74p.
ROBERTS, C. M. (1998). “Sources, sinks, and the design of marine reserve networks.” Fisheries
23(7): 16-19.
ROBERTS, C. M. and N. V. C. POLUNIN (1993). “Marine reserves: simple solutions to managing
complex fisheries?” Ambio 22(6): 363-368.
ROWLEY, R. J. (1994). “Case studies and reviews: Marine reserves in fisheries management.”
Aquatic Conservation: marine and freshwater ecosystems 4: 233-254.
SANCHIRICO, J. N. and J. E. WILEN (1999). “Bioeconomics of spatial exploitation in a patchy
environment.” Journal of Environmental Economics and Management 37: 129-150.
SANCHIRICO, J. N. and J. WILEN (2000b). Spatial dynamics of overexploitation : a metapopulation
approach, Discussion paper 00-25, Resources for the Future.
SANCHIRICO, J. N. and J. E. WILEN (2000a). The impacts of marines reserves on limited-entry
fisheries. Discussion paper n°00-34. Washington DC, Resources for the Future.
SHACKELL, N. and WILLISON, M.J.H. (eds) (1995) Marine protected areas and sustainable
fisheries. Science and Management of Protected Areas Association, Wolfville, Nova Scotia, Canada,
300 p.
SLADEK NOWLIS, J. and C. M. ROBERTS (1999). “Fisheries benefits and optimal design of
marine reserve.” Fishery Bulletin 97(3): 604-616.
SOMERTON, D. A. and J. JUNE (1984). “A cost-benefit method for determining optimum closed
fishing areas to reduce the trawl catch of prohibited species.” Canadian Journal of Fisheries and
Aquatic Sciences 41: 93-98.
SUMAILA, U. R. (1998). “Protected marine reserves as fisheries management tools: a bioeconomic
analysis.” Fisheries Research 37(1-3): 287-296.
SUPRIATNA, A. K. et H. P. POSSINGHAM (1999). “Harvesting a two-patch predator-prey
metapopulation.” Natural Resource Modeling 12(4): 481-498.
13
DRAFT – Please do not copy or cite without permission of the authors.
THOMSON, C. (1998). Evaluating marine harvest refugia: an economic perspective. in Marine
harvest refugia for West Coast Rockfish. M.M. YOKLAVICH, NOAA, Technical Memorandum
NMFS, N°255.
TUCK, G.-N. and H.-P. POSSINGHAM (1994). “Optimal harvesting strategies for a
metapopulation.” Bulletin of Mathematical Biology 56(1): 107-127.
WHITMARSH, D., C. JAMES, et al. (2000). Economic effects of marine protected areas on smallscall fisheries: a case study of the trawl ban in the Gulf of Castellammare, Sicily. 10th Biennal
Conference of IIFET, 10-14 July, Corvallis, Oregon, USA.
WILSON, J., B. LOW, et al. (1999). “Scale misperceptions and the spatial dynamics of the socialecological system.” Ecological Economics 31(2): 243-257.
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