A framework for incorporating climate regime

shifts into the management of marine resources

J . R. KI NG & G. A. MCF ARL ANE
Pacic Biological Station, Fisheries and Oceans Canada, Nanaimo, British Columbia, Canada
Abstract It is possible to use an ecosystem-based management approach to incorporate knowledge of climate
regime impacts on ecosystem productivity to manage shery resources. To do so, it requires the development of a
coherent framework that can be built using existing stock assessment and management activities: ecosystem
assessment, risk analyses, adaptive management and reference points. This paper builds such a framework and
uses two population simulations to illustrate the benets and tradeos of variable regime-specic harvest rates.
The framework does not require prediction of regime shifts, but assumes that detection can occur soon after one
has happened. As such, decisions do not need to be coincident to regime shifts, but can be delayed by an
appropriate period of time that is linked to a species life history, i.e. age of maturity or recruitment. Fisheries
scientists should provide harvest recommendations that reect a range of levels of risk to the stock under dierent
assumptions of productivity. Coupling ecosystem assessment with ecosystem-based management would allow
managers to select appropriate regime-specic harvest rates.
KEYWORDS : ecosystem assessment, ecosystem-based management, regime shifts.
Introduction
While there may be debate on the magnitude of the
decline of the worlds marine resources or the magni-
tude of overexploitation by existing sheries [Pauly,
Christensen, Dalsgaard, Froese & Torres 1998;
Jackson, Kirby, Berger, Bjorndal et al. 2001; Fisheries
and Agriculture Organization of the United Nations
(FAO) 2002; Hilborn, Branch, Ernst, Magnusson,
Minte-Vera, Scheurell & Valero 2003; Myers & Worm
2003; Walters 2003], the vast majority of sheries
scientists agree that some of the worlds sheries have
collapsed, many are overexploited and most are
intensely exploited. The concerns regarding over-
exploitation of marine resources extend beyond the
impacts of sheries on sh populations and have begun
to include the interactions between sh populations
(and other ecosystem interactions) and the overall
impact of climate variability on sh populations
(Beamish 1995; McFarlane, King & Beamish 2000;
Bakun 2004). Admittedly there has been disparity in
views regarding the relative importance of climate
variability and of shing mortality on the regulation
of sh populations; however, accounting for either
process (or both), in conjunction with species interac-
tions, falls within the frameworks of ecosystem assess-
ment and ecosystem-based sheries management.
Although often used interchangeably, the terms
ecosystem assessment and ecosystem-based sheries
management refer to two distinct approaches in stock
assessment and management. Ecosystem assessment
typically refers to monitoring climateocean indices
and indicator species to detect ecosystem changes, i.e.
assessing the state of the environment. Ecosystem-
based sheries management attempts to account for
ecosystem processes when formulating sheries man-
agement advice (Sissenwine & Murawski 2004). This
approach would incorporate knowledge of the state of
the ecosystem (i.e. the ecosystem assessment) into
stock assessments when making management decisions
[e.g. King, McFarlane & Beamish 2001; Boldt 2004;
Department of Fisheries and Oceans Canada (DFO)
2004]. Ecosystem-based sheries management can also
encompass the consideration of shing impacts on
non-target species (components of the ecosystem) and
habitats. All in all, the term has come to represent a
more holistic approach to stock assessment (consider-
ing interactions between physical and biological
Correspondence: Dr Jacquelynne King, Pacic Biological Station, Fisheries and Oceans Canada, Nanaimo, BC, Canada V9T 6N7 (e-mail:
kingjac@pac.dfo-mpo.gc.ca)
Fisheries Management and Ecology, 2006, 13, 93102
2006 Blackwell Publishing Ltd 93
ecosystem components) and sheries management
(recognising that sheries comprised people and sh,
and have considerable impacts on ecosystems) (see
Browman & Stergiou (2004) for a collection of
perspectives on ecosystem-based approaches).
The underlying paradigm for ecosystem-based sh-
eries management is that sh populations may be
impacted by exploitation, but the fundamental
mechanisms for sh population dynamics are changes
in ecosystems, including climateocean impacts and
trophic level interactions. Marine shes have evolved
life history strategies to cope with variability in their
environment, ranging from short-lived species with
highly variable stock dynamics that respond immedi-
ately to changes in their environment to extremely
long-lived species whose population dynamics are
mainly stable (King & McFarlane 2003). Short-lived
species (opportunistic strategists) have a short genera-
tion time that compensates for relatively low fecundity
and therefore maximises their intrinsic rate of popula-
tion growth. Long-lived species (periodic strategists)
are typically highly fecund and have a long period
of reproductive potential (i.e. greater than 20 years),
which allows them to take immediate advantage of
changes to more productive periods, through increased
year class success. Longevity allows a species to
persist through prolonged periods of poor productivity
(Leaman & Beamish 1984; McFarlane & Beamish
1986).
Climate variability as it impacts on marine ecosys-
tems occurs on three time scales: interannual (El Nin o-
La Nin a southern oscillation (ENSO) events), decadal
(regimes and regime shifts) and long-term (global
warming trends). For sh stocks, the most important
scale of variability from a sheries management point
of view is the decadal-scale variability. Typically,
ENSO events, while they can aect sh survival,
mainly impact the distribution of sh populations and
have short-lived implications for year class success. At
the opposite end of the spectrum, long-term global
warming impacts are yet to be clearly dened and will
require the long-range planning of agencies to deal
with the likely irreversible consequences of either a loss
of a sh species or the introduction of a new species
within their jurisdiction. Marine resource management
can deal with the decadal-scale variability of climate
ocean regimes. A regime is a period of a decade or
more in which the state, or characteristic behaviour, of
the climate or ocean system is steady. Year-to-year
dierences may exist, but overall the state of the
system varies around a persistent baseline. A regime
shift is rapid, usually occurring within a year, and is a
substantial change from one regime period to another.
Scientic literature reports regime shifts in the North
Pacic occurring in 1925, 1947, 1977, 1989 and 1998
(Mantua, Hare, Zhang, Wallace & Francis 1997; Hare
& Mantua 2000; McFarlane et al. 2000; Minobe 2000;
Schwing & Moore 2000; Bond, Overland, Spillane &
Stabeno 2003; King 2005).
Numerous studies have illustrated that climate
ocean regimes are associated with uctuation in sh
abundance and population dynamics (Kawasaki &
Omori 1986; Beamish & Bouillon 1993; Polovina 1996;
Mantua et al. 1997; McFarlane et al. 2000; King 2005).
However, to date few attempts have been made to
incorporate the dynamics of regimes into the manage-
ment of sh populations. The only example might be
groundsh stock assessments conducted for the Gulf of
Alaska (Livingston, Aydin, Boldt, Ianelli & Jurado-
Molina 2005). Incorporating climate regime impacts
into marine resource management has been impeded by
a lack of a coherent framework despite management
tools to do so successfully having been developed and
reported in the sheries scientic literature. In this
paper an applicable management tools and the outline
of a suitable management framework that allows for
the management of marine resources given climate
regime shift impacts are reviewed. The application of
the proposed framework is illustrated for two theo-
retical sh populations that encompass two life history
strategists: short-lived, (opportunistic) strategists and
long-lived, (periodic) strategists.
Building an ecosystem-based sheries
management framework
Science and management activities are already avail-
able for incorporating the inherent dynamics of sh
populations as they respond to climateocean regime
shifts and subsequent changes in ecosystem productiv-
ity. These activities need to be organised in a decision-
based framework that outlines a priori the responses in
management that would result from indications that a
regime shift has occurred. An example of such a
framework is outlined in Figure 1, together with the
likely actions taken under dierent regime scenarios.
Monitoring of basin-scale climate and ocean indices
and regional ecosystem parameters (ecosystem assess-
ment) would provide the means for the detection of
climateocean regime shifts and changes in productiv-
ity. The framework outlined here does not require the
prediction of a regime-shift, but would require meth-
ods of regime shift detection that are currently being
developed (Rodionov & Overland 2005). The response
time of sheries management can mirror the response
time of biota to changes in productivity as it impacts
J. R. KI NG & G. A. MCFARLANE 94
2006 Blackwell Publishing Ltd, Fisheries Management and Ecology, 2006, 13, 93102
the shery of interest. Stock assessment and risk
analyses approaches can incorporate assumptions
about regime impacts on sh recruitment and can
provide sheries managers a range of harvest rates
with varying levels of risk to the population from
which to select regime-specic harvest rates. Reference
points provide the check system and stop mechanism
to help avoid overexploitation.
Ecosystem assessment
To incorporate the climateocean dynamics into
marine resource management successfully, agencies
need to identify and document the state of the
environment to recognise that signicant changes, or
shifts, have occurred. There are global or basin-wide
climateocean indices that provide indication of large-
scale climate regime shifts, but the impacts of these
shifts may differ on a regional scale. As such, it is
necessary to monitor both physical and biological
indices on a regional scale in conjunction with basin-
wide indices. For example, King et al. (2001) used a
matrix of climatological and oceanographic indices for
the North Pacic, and regional environmental and
biological indices for the west coast of Vancouver
Island, British Columbia, as a report card of produc-
tivity during specic regimes to augment stock assess-
ment and management eorts of sablesh,
Anoplopoma mbria Pallas. The report card approach
was intended to provide a general indication of the
overall state of the ecosystem and was built upon
conceptual mechanisms linking lower trophic produc-
tivity to year class success. King et al. (2001) is an
example of ecosystem assessment linked to a specic
target species, but there are more broad-scale exam-
ples, such as the Ecosystem Considerations Appendix
that is now routinely available with annual Stock
Assessment and Fisheries Evaluation reports produced
by the Alaska Fisheries Science Center (see, for
example, Boldt 2004). The Ecosystem Considerations
Appendix provides an extensive overview of North
Pacic climate and regime shift indicators along with
environmental and biological ecosystem status indica-
tors for the Gulf of Alaska, Aleutian Islands and the
Eastern Bering Sea. Information on such variables as
the physical environment, nutrients and productivity,
zooplankton, forage and benthic sh, marine mam-
mals, sea birds and community indices are provided as
a source of overall ecosystem status trends that is
intended to augment target species assessments.
Timeframes for responses
Fisheries are typically conducted on mature sh, such
that the age of recruitment to the shery generally
corresponds to the age of maturity. Age of maturity
Reference points
Is the stock above critical
spawning biomass?
Risk analysis
Select a regime-
specific harvest rate
based on acceptable
level of risk and a
productivity
assumption
Low
risk
High
risk
Low
productivity HR1 HR2
High
productivity HR3 HR4
Variable harvest
Rates across regimes
Close
fishery
Maintain
harvest rate
Stock
assessment
Estimate of
spawning stock
Yes No
No Yes
Adaptive management
Ecosystem assessment
Has a regime shift occurred?
Figure 1. Conceptual decision-rule framework using existing science and management actions available to conduct ecosystem-based sheries
management which includes ecosystem monitoring or assessment for the detection of regime shifts and sets regime-specic harvest rates (HR).
REGI MES AND MARI NE RESOURCE MANAGEMENT 95
2006 Blackwell Publishing Ltd, Fisheries Management and Ecology, 2006, 13, 93102
can be used as an indication of the length of time that
sheries managers have to respond to a shift in
productivity. For example, Pacic cod, Gadus macro-
cephalus Tilesius, matures and recruits to the shery at
age 2+ years, so a shift in productivity that affects
year class success would have impacts on the biomass
available to the shery in 3 years (i.e. a strong year
class will appear as a large increase in biomass for the
shery within 3 years). Conversely, a large year class in
a rocksh species (Sebastes spp.) would not appear as
an increase in biomass to the shery for 810 years
(depending on the species). The response time of
management actions to regime shifts may be lagged by
a correspondence to the age of recruitment of the
species of interest. For example, sheries management
can use decision rules to outline actions that will be
taken when there are indications that a regime shift has
occurred. The response time for the actions resulting
from these decision rules should be linked to the
species lifespan, age of maturity and rate of produc-
tion.
Stock assessment and risk analysis
Fisheries scientists should provide harvest recommen-
dations that reect a range of risks (low to high) to the
stock under different assumptions of productivity or
recruitment (e.g. low, medium and high year class
success). A low level of risk would guarantee a minimal
impact of shing on the stock, but would be associated
with a signicant economic impact on the industry. A
high risk level, could increase conservation concerns
for a stock but may be chosen for social, economic or
political considerations. Differing levels of productiv-
ity appear to be decadal in nature and correspond to
regimes; changes in productivity coincide with regime
shifts (Hare & Mantua 2000; McFarlane et al. 2000;
Hollowed, Hare & Wooster 2001). The ecosystem
assessment should provide an indication to the stock
assessment scientists, and to the managers, which
productivity scenario is most appropriate. Scientic
advice will remain only one of the myriad of factors
that managers use to make decisions. Managers will
need to consider economic, political and social factors
when selecting the appropriate level of risk to the stock
that they are willing to accept and select a regime-
specic harvest rate accordingly.
Reference points
A number of countries have adapted the philosophical
precautionary approach to managing marine
resources. This approach is embodied in a number of
international agreements, such as the Rio Declaration
on Environment and Development [United Nations
Conference on Environment and Development
(UNCED) 1992], the FAO Code of Conduct for
Responsible Fisheries (FAO 1995) and the United
Nations Convention on the Law of the Sea [United
Nations (UN) 1995] provisions that are related to the
conservation and management of highly migratory sh
stocks and sh stocks with distributions that span
across political boundaries. The primary mechanism
for applying the precautionary approach is the use of
limit reference points. Limit reference points take into
account the reproductive capacity of the population
and are often expressed in terms of stock spawning
biomass or numbers of mature individuals (Rivard
1999). Most research on limit reference points have
focused on target species for which there is adequate
time series of sheries statistics, abundance indices and
biological data. Eort on developing limit reference
points for non-target species would be required in an
ecosystem-based approach and could include synoptic
data collected on age composition, research catch rates
or bycatch in monitored sheries.
Variable harvest rates
Simulation studies that have investigated harvest
strategies for sheries that are impacted by climate-
induced shifts in productivity, similar to regime shifts,
have reached different conclusions. Walters & Parma
(1996) concluded that overall, a constant rate harvest
strategy performs well; while other studies suggest that
a regime-specic harvest rate strategy is optimum
(Spencer 1997; Peterman, Pyper & Grout 2000).
MacCall (2002) suggested that for short-lived species
(dened as a sh with a lifespan less than 30 years), a
regime-specic harvest rate was optimum for main-
taining high yield and low variation in spawning stock
biomass. The harvest rate did not need to change with
the regime shift year, but could be lagged by a few
years. Conversely, for a long-lived species (lifespan
greater than 30 years), a constant harvest rate was
more appropriate (MacCall 2002).
Polovina (2005) suggested that for sheries with
regime impacts on productivity, if a constant harvest
rate strategy is used, the harvest rate must be well
below traditional benchmarks (i.e. approximately 10%
of the exploitable biomass), to avoid overshing during
low productivity regimes. The obvious trade-o is that
a low harvest rate results in low overall yield from the
stock, particularly during high productivity regimes.
Polovina (2005) noted that a regime-specic harvest
rate strategy would be benecial by increasing yield in
J. R. KI NG & G. A. MCFARLANE 96
2006 Blackwell Publishing Ltd, Fisheries Management and Ecology, 2006, 13, 93102
high productivity regimes, but might result in shery
closures during regimes of low productivity. Within the
proposed framework here, sheries scientists and
managers would use the ecosystem assessment to
detect that a regime shift has occurred (this detection
can be lagged by several years), and from the stock
assessment advice and risk analyses, sheries managers
could select a harvest rate that would be applicable to
the upcoming productivity regime.
Illustrating the benets and trade-offs of regime-
specic harvest rates
To illustrate the benets and trade-offs of using a
response time that corresponds to age of recruitment
and of using regime-specic harvest rates, two hypo-
thetical populations were simulated: a short-lived
species and a long-lived species (Table 1). Both pop-
ulations were simulated with a Beverton-Holt-based
age-structured model, with regime impacts added as a
multiplicative eect on the slope at the origin:
R
i

1
ae
i
b=P
i1

;
where R is the number of young in year i, P are the
number of spawners at the end of the previous year
after any harvest (expressed as number sh), e is the
regime effect assigned to year i and a and b are
parameters (Table 1). Three, 20-year regime periods
were modelled, with the rst 20 years as a period of
good productivity (regime 1), the second 20 years as a
period of poor productivity (regime 2) and the nal
20 years as a period of moderate productivity (regime
3). Values of e were selected such that the resultant
unshed biomass approximated 50% in regime 2 and
75% in regime 3 of the regime 1 biomass (Table 1).
Two sets of harvest scenarios were used and the total
harvest (tonnes) was calculated over the 60 years for
each scenario. The rst group of scenarios used a
constant harvest rate (F) over the 60 years (irrespective
of changes in productivity) set to F annual mortality
(M); F 0.5M and F 0.25M. The second set used
regime-specic harvest rates with scenarios varying by
the timing in which the harvest rate was switched:
coincidental with the regime shift year, delayed until
age of 50% maturity (i.e. 3 or 5 years for short-lived or
long-lived respectively) and delayed until twice the age
of 50% maturity. One group of regime-specic harvest
rate scenarios used F M, F 0.5M and F M for
regime 1, regime 2 and regime 3 respectively. These
represent typical harvest rates proposed in sheries
management. The other regime-specic harvest rate
scenarios used F M, F 0.25M and F 0.5M for
regime 1, regime 2 and regime 3 respectively, to reect
the relative levels of productivity for each respective
regime (i.e. high, low and moderate).
The maximum number of spawners observed over
the 60 years without any harvest was used to set the
criteria for shery closures and conservation concerns.
The 25% and 30% levels of this maximum were used
as representatives of critical spawning biomass and a
level for which there would be conservation concerns
respectively. These levels are currently used by North
American management agencies (Fisheries and Oceans
Canada; National Marine Fisheries Service) as mini-
mum spawning biomass reference points. When run-
ning the harvest scenarios, the harvest rate would be
set to zero if the spawning stock (numbers of sh) fell
below the critical spawning biomass. The number of
years that the shery was closed and the total number
of years in which there was a conservation concern
were totalled for all 60 years.
Short-lived species
The simulated unshed population of the short-lived
species responded to regime shift changes in year class
success (i.e. productivity) quickly and the resultant
lower or higher spawning stock size stabilised within
5 years (Fig. 2a). As expected, the spawning stock was
depleted more quickly and to a lower level as the
constant harvest rate increased from F 0.25M to M
(Fig. 2a). Conversely, the rebuilding rate and level to
which that the spawning stock rebuilt during the third
regime when year class success improved was greater
for the lower constant harvest rate (F 0.25M)
scenario (Fig. 2a). In the simulation, the aggressive
harvest rate of F M translated into a failure of the
spawning stock to rebuild, and the spawning stock
remained close to the level that was dened as the
critical spawning stock size (Fig. 2a).
By denition, the total yield decreased with decreas-
ing constant harvest rate, such that the total yield with
Table 1. Parameters used in the Beverton-Holt age-structured
models for the short-lived and long-lived species
Parameters Short-lived species Long-lived species
Maximum age 10 50
Age at 50% maturity 3 5
Annual mortality 0.4 0.1
Maximum size (kg) 0.3 3
A 0.00000017 0.000001
B 0.02 0.001
Regime 1 eect, e 1 1
Regime 2 eect, e 1.7 3
Regime 3 eect, e 1.2 1.5
REGI MES AND MARI NE RESOURCE MANAGEMENT 97
2006 Blackwell Publishing Ltd, Fisheries Management and Ecology, 2006, 13, 93102
F 0.25M was only 30% of the total yield achieved
with F M (Fig. 3a). However, the benet of a lower
constant harvest rate in this modelled population was
fewer or no shery closures or years with a conserva-
tion concern (Fig. 3b). The aggressive constant harvest
rate (F M) resulted in 8 years of closure with an
additional 21 years in which there would be a conser-
vation concern, which means 48% of the time the
spawning stock is near or below the critical spawning
stock size that was set as the limit reference point
(Fig. 3b). Overall, lower constant harvest rates of F
0.5M or F 0.25M resulted in no shery closures and
no years with a conservation concern, but there was a
substantial decrease in the total yield obtained
(Fig. 3a, b).
The regime-specic harvest rate scenario with F
M, F 0.5M and F M allowed for aggressive
harvesting in periods of good productivity, producing
comparable total yields to the constant F M
scenario and higher yields than the constant F
0.5M or F 0.25M scenarios (Fig. 3a). They pro-
duced fewer years with a shery closure (12 years) or
a conservation concern than the constant F M
scenario (Fig. 3b). There was no advantage in switch-
ing the harvest rates exactly on the year of the regime
shift and a delay to the age of recruitment in
switching harvest rates did not increase the years
with a shery closure but reduced the number of
years with conservation concerns (Fig. 3b). Delaying
the switch in harvest rates by twice the age of
recruitment increased the number of years in which
there was a shery closure (Fig. 3b). For these
regime-specic harvest rate scenarios, the stock failed
to rebuild during regime 3, when year class success
improved (Fig. 2b). The benet of increased total
yield was traded-o by the years with shery closures
or conservation concerns and the lack of rebuilding
when recruitment improves.
These trade-offs were minimised with the selection
of regime-specic harvest rates that reected the
relative productivity of the different regimes (F M,
F 0.25M and F 0.5M). The total yield obtained
under this harvest rate scenario was 15% higher than
that obtained with F 0.5 M (Fig. 3a) and there were
no years with shery closures or conservation concerns
unless the change in harvest rate is delayed by twice the
age of recruitment (Fig. 3c). More importantly, the
spawning stock size was higher during the low
productivity regime than the spawning stock size
maintained with a constant F 0.50M harvest rate,
and the spawning stock was able to rebuild to
comparable levels during regime 3, when productivity
improved (Fig. 2c).
(a)
0
1
2
3
4
5
0 5 10 15 20 25 30 35 40 45 50 55 60
Year
(b)
0
1
2
3
4
5
0 5 10 15 20 25 30 35 40 45 50 55 60
Year
S
p
a
w
n
i
n
g

s
t
o
c
k

(
m
i
l
l
i
o
n
s

o
f

f
i
s
h
)
S
p
a
w
n
i
n
g

s
t
o
c
k

(
m
i
l
l
i
o
n
s

o
f

f
i
s
h
)
Year
(c)
S
p
a
w
n
i
n
g

s
t
o
c
k

(
m
i
l
l
i
o
n
s

o
f

f
i
s
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)
0
1
2
3
4
5
0 5 10 15 20 25 30 35 40 45 50 55 60
Figure 2. (a) Spawning stock (millions of sh) from a Beverton-Holt
age-structured model for the short-lived population with no harvest
(solid line), with harvest rate (F) equal to annual mortality (M) (dotted
line), with F 0.5M (dotted and dashed line) and with F 0.25M
(dashed line). (b) Spawning stock (millions of sh) from the same
model, but with regime-specic harvest rates (F M, F 0.5M and
F M) where the switch in harvest rates coincides with the regime shift
year (solid line), is delayed by age of 50% maturity (3 years, dashed
line) or is delayed by twice the age of 50% maturity (dotted line). (c)
Spawning stock (millions of sh) with regime-specic harvest rates
(F M, F 0.25M and F 0.5M) where the switch in harvest rates
coincides with the regime shift year (solid line), is delayed by age of 50%
maturity (3 years, dashed line) or is delayed by twice the age of 50%
maturity (dotted line); the spawning stock size for the constant harvest
rate F 0.5M (dotted and dashed line) is redisplayed for comparison.
In all plots, the horizontal line indicates the assigned critical spawning
stock size below which no harvest is permitted. In all model simulations,
the rst 20-year regime period is high productivity, the second 20-year
regime period is low productivity and the third regime period is mod-
erate productivity.
J. R. KI NG & G. A. MCFARLANE 98
2006 Blackwell Publishing Ltd, Fisheries Management and Ecology, 2006, 13, 93102
Long-lived species
The spawning stock of the unshed population began
to respond to the regime shifts 5 years (age to 50%
maturity) after they occurred (Fig. 4a). The decrease
or increase of spawning stock size in response to
regime shifts to low or moderate productivity contin-
ued throughout the regime period and did not stabilise
(Fig. 4a). The higher the rate of harvest in the constant
harvest rate scenarios, the greater the rate and level of
depletion during the low productivity regime, and the
lower the level of rebuilding in the moderate produc-
tivity regime (Fig. 4a). The benet of higher total yield
obtained with an aggressive harvest rate (F M) was
offset by 10 years with shery closures and an addi-
tional 18 years with conservation concerns (Fig. 5a, b).
Even the F 0.5M constant harvest rate scenario
resulted in years with shery closures and with
conservation concerns (Fig. 5b). Only the lowest har-
vest rate (F 0.25M) did not result in these draw-
backs (Fig. 5b), but the total yield was approximately
60% lower than the total yield obtained from F M
(Fig. 5a).
The regime-specic harvest rate scenario with F
M, F 0.5M and F M produced high total yields,
comparable with the constant harvest rate scenario
F M (Fig. 5a). Switching harvest rates the same year
as a regime shift, has an immediate impact on
spawning stock size when the year class success goes
from high to low; the immediate reduction in harvest
rate results in a 5-year increase in spawning stock size
as a pulse of good recruitment from the previous
regime enters the shery (Fig. 4b). Delaying the switch
to regime-specic harvest rates by the age of recruit-
ment or twice this age (5 and 10 years respectively), did
not alter the ability of the spawning stock to rebuild
(Fig. 4b) or greatly alter the number of years with
shery closures or with conservation concerns
(Fig. 4b). Unlike the short-lived species, when produc-
tivity improved (regime 3), the spawning stock exhib-
ited moderate rebuilding (Fig. 4b). This is a function of
the prolonged reproductive lifespan of a long-lived
species and the benet of having many age classes. It
also highlights the benet of maintaining a robust age
structure and conserving older sh.
As with the short-lived species, the regime-specic
harvest rate scenario with F M, F 0.25M and
F 0.5M reduced the number of years with a shery
closure (Fig. 5c) and produced slightly higher overall
yields than the constant F 0.5M or F 0.25M
harvest rates (Fig. 5a). This scenario allowed aggres-
sive harvesting during the high productivity regime
(regime 1), but the spawning stock size by the end of
(a)
0
2
4
6
8
10
12
14
16
18
F = M F= 0.5M F= 0.25M No lag 3-year lag 6-year lag
3-year lag 6-year lag
3-year lag 6-year lag
Model scenario
Y
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d

(
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n
d
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o
f

t
o
n
n
e
s
)

(b)
F = M F = 0.5M F= 0.25M No lag
Model scenario
0
5
10
15
20
25
N
u
m
b
e
r

o
f

y
e
a
r
s
(c)
F = M F = 0.5M F =0.25M No lag
Model scenario
0
5
10
15
20
25
N
u
m
b
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r

o
f

y
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Figure 3. (a) The total harvest (thousands of tonnes) for the various
harvest rate scenarios for the short-lived species population; solid bars
are the constant harvest rate scenarios and the F M; F 0.5M; F
M regime-specic harvest rate scenarios; open bars are the F M;
F 0.25M; F 0.5M regime-specic harvest rate scenarios. (b) The
number of years that the shery was closed (solid bars) or in which there
was a conservation concern (open bars) as dened by 25% and 30% of
the maximum unshed spawning stock respectively. Six harvest rate
scenarios are presented: constant harvest rates (F) for 60 years, F equal
to annual natural mortality (M) or F 0.5M or F 0.25M; and
regime-specic harvest rates (F M; F 0.5M; F M) in which the
switch in harvest rates coincides with the regime shift year (no lag), is
delayed by age of 50% maturity (3-year lag) or by twice the age of 50%
maturity (6-year lag). (c) Similar to (b) with the constant harvest rate
scenarios redisplayed for comparison with the regime-specic harvest
rates (F M; F 0.25M; F 0.5M).
REGI MES AND MARI NE RESOURCE MANAGEMENT 99
2006 Blackwell Publishing Ltd, Fisheries Management and Ecology, 2006, 13, 93102
(a)
0
1
2
3
4
5
6
1 6 11 16 21 26 31 36 41 46 51 56 61
Year
S
p
a
w
n
i
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g

s
t
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c
k

(
m
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l
i
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o
f

f
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h
)
(b)
0
1
2
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6
1 6 11 16 21 26 31 36 41 46 51 56 61
Year
S
p
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k

(
m
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f
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)
(c)
0
1
2
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6
1 6 11 16 21 26 31 36 41 46 51 56 61
Year
S
p
a
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(
m
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h
)
Figure 4. (a) Spawning stock (millions of sh) from a Beverton-Holt
age-structured model for the long-lived population with no harvest
(solid line), with harvest rate (F) equal to annual mortality (M) (dotted
line), with F 0.5M (dotted and dashed line) and with F 0.25M
(dashed line). (b) Spawning stock (millions of sh) from the same
model, but with regime-specic harvest rates (F M, F 0.5M and
F M) where the switch in harvest rates coincides with the regime shift
year (solid line), is delayed by age of 50% maturity (5 years, dashed
line) or is delayed by twice the age of 50% maturity (dotted line). (c)
Spawning stock (millions of sh) with regime-specic harvest rates
(F M, F 0.25M and F 0.5M) where the switch in harvest rates
coincides with the regime shift year (solid line), is delayed by age of 50%
maturity (5 years, dashed line) or is delayed by twice the age of 50%
maturity (dotted line); the spawning stock size for the constant harvest
rate F 0.5M (dotted and dashed line) is redisplayed for comparison.
In all plots, the horizontal line indicates the assigned critical spawning
stock size below which no harvest is permitted. In all model simulations,
the rst 20-year regime period is high productivity, the second 20-year
regime period is low productivity and the third regime period is mod-
erate productivity.
(a)
0
5
10
15
20
25
30
35
F=M F=0.5M F = 0.25M No lag
Model scenario
Y
i
e
l
d

(
t
h
o
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s
a
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d
s

o
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s
)
(b)
F=M F=0.5M F =0.25M No lag
Model scenario
0
2
4
6
8
10
12
14
16
18
20
N
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b
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r

o
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y
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r
s
(c)
F=M F =0.5M F =0.25M No lag 5-year lag 10-year lag
5-year lag 10-year lag
5-year lag 10-year lag
Model scenario
0
2
4
6
8
10
12
14
16
18
20
N
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m
b
e
r

o
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Figure 5. (a) The total harvest (thousands of tonnes) for the various
harvest rate scenarios for the long-lived species population; solid bars
are the constant harvest rate scenarios and the F M; F 0.5M; F
M regime-specic harvest rate scenarios; open bars are the F M;
F 0.25M; F 0.5M regime-specic harvest rate scenarios. (b) The
number of years that the shery was closed (solid bars) or in which there
was a conservation concern (open bars) as dened by 25% and 30% of
the maximum unshed spawning stock respectively. Six harvest rate
scenarios are presented: constant harvest rates (F) for 60 years, F equal
to annual natural mortality (M) or F 0.5M or F 0.25M; and
regime-specic harvest rates (F M; F 0.5M; F M) in which the
switch in harvest rates coincides with the regime shift year (no lag), is
delayed by age of 50% maturity (5-year lag) or by twice the age of 50%
maturity (10-year lag). (c) Similar to (b) with the constant harvest rate
scenarios redisplayed for comparison with the regime-specic harvest
rates (F M; F 0.25M; F 0.5M).
J. R. KI NG & G. A. MCFARLANE 100
2006 Blackwell Publishing Ltd, Fisheries Management and Ecology, 2006, 13, 93102
the low productivity regime (regime 2) was higher than
that under a traditional constant harvest rate F
0.5M, and the rebuilding of the spawning stock when
productivity improves (regime 3) was similar (Fig. 4c).
Recommendations
Agencies interested in implementing ecosystem-based
sheries management can take advantage of the
existing sheries science and management activities
and build frameworks similar to the one outlined. In
doing so, ecosystem assessments will become part of
the larger ecosystem-based sheries management and
can be used to detect changes in productivity related to
regime shifts. These frameworks need to specify
explicit decision rules and subsequent actions to be
taken in response to preliminary indications that a
regime shift has occurred. Stock assessment advice
should provide an indication of the likely consequences
to stock viability of alternate harvest strategies under
various recruitment assumptions. Decision rules must
use reference points such as critical spawning biomass.
In the modelling exercises, the regime-specic harvest
rates that reected the relative levels of productivity by
regime (i.e. high, low, moderate) produced the best
balance between benets (high yield) and trade-offs
(shery closures) and overall allowed for rebuilding of
the spawning stock when productivity improved in a
similar timeframe to a similar level to that produced
from the traditional constant harvest rate F 0.5M.
For long-lived species, this recommendation is con-
trary to MacCall (2002), who suggested a constant
harvest rate was more appropriate. However, MacCall
(2002) focused on optimising yield and minimising
timeframes for rebuilding. The recommendations made
here may reect the additional consideration of
avoiding years with shery closures.
Given the dramatic impacts that regime shifts have
on productivity, constant harvest rates that are typic-
ally employed (e.g. F M) cannot maintain sustain-
able stock or sheries when productivity is greatly
reduced. It is possible to use constant harvest rates
across productivity regimes, but these harvest rates
must be very low (e.g. F 0.25M), as suggested by
Polovina (2005). These very low harvest rates are
partnered with the trade-o of much reduced yield.
The resulting loss of yield might have signicant
economic and social impacts on industry. Regime-
specic harvest rates can be delayed in implementation
by the age of maturity with very little impact on total
harvest or the number of years with shery closures.
Even if a constant harvest rate was applied across
regimes, the ecosystem assessment would still be
required to provide information on recruitment to
estimate exploitable biomass. Given the benets of
these approaches, and because the activities already
exist to build suitable frameworks, agencies can now
move forward with their mandates to implement
ecosystem-based sheries management and begin to
incorporate regime shift impacts on sh productivity
into the way they assess and manage marine resources.
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