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Regime shift in marine ecosystems and implications for fisheries

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Rev Fish Biol Fisheries (2009) 19:177–191
DOI 10.1007/s11160-008-9096-8
Regime shift in marine ecosystems and implications
for fisheries management, a review
Yan Jiao
Received: 29 May 2008 / Accepted: 12 September 2008 / Published online: 19 October 2008
Ó Springer Science+Business Media B.V. 2008
Abstract The concept of regime shift in marine
ecosystems has arisen in recent years. This study
reviews the origin, scientific meaning, driving forces
of marine ecosystem regime shrifts. Driving forces
include climate-ocean oscillation, fishing, introduced
species, river flow, eutrophication, disease and pol-
lution. Several controversial questions are discussed:
whether regime shift is just a climate oscillation
noise, the difficulties of separating driving forces, and
whether the impacts of ecosystem regime shift are
necessarily bad or good. The methods used to model
regime shifts in ecosystems are reviewed. These
methods include two types of models: general
circulation model and historical description. The
implications of regime shift in fisheries are consid-
ered, including examples of impacts, consequences of
not considering regime shift, and difficulties in
incorporating regime shift into the current stock
assessment models. Possible future research needed
to understand and model regime shifts is discussed,
including strategies for fisheries management.
Keywords Regime shift
Marine ecosystem

Fisheries
Red noise
General circulation model
Y. Jiao (&)
Department of Fisheries and Wildlife Sciences, Virginia
Polytechnic Institute and State University, Blacksburg,
VA 24061-0321, USA
e-mail: yjiao@vt.edu
Regime shift defined
Until recently the word ‘‘regime’’ was not in the
vocabulary of many fisheries biologists. Even today,
scientists are not entirely sure how to define regimes.
Regime in the dictionary has two meanings: a regular
pattern of occurrence or action (as of seasonal
rainfall); and the characteristic behavior or orderly
procedure of a natural phenomenon or process
(Merriam-Webster’s Collegiate Dictionary 1998).
Isaacs (1976) introduced to fishery scientists the
concept of a number of possible ‘locked-in persistent
episodes’, or ‘regimes’, in climate, oceanography and
biological systems (Steele 1996; Beamish et al. 1999,
2000). He suggested that these regimes are connected
by a number of abrupt discontinuities. Regimes are
large, linked climate-ocean ecosystems that shift over
10–30-year periods (Beamish and Mahnken 1999). A
regime shift is a change in the mean of a data series
and the shift is completed within several months to a
year (Beamish et al. 2000).
Ecosystems are dynamic due to chaotic distur-
bance inside and outside of the community. Though
most evidence of shifts is provided by economic
species, the term ‘‘regime shift’’ in ecosystems
suggests that changes are causally connected and
can be linked to other changes in non-commercial
species, such as planktonic invertebrates. Climate
changes alter abiotic environmental factors and
impact biotic diversity and richness. Fluctuations or
successions of the community can occur, as in
123
178
the sardine/anchovy fluctuation in the eastern Pacific,
zooplankton abundance changes, fish community
changes in the northeast Pacific, and the comple-
mentary trend of fish and shellfish landings in
the northwest Atlantic Ocean (Ebbesmeyer et al.
1991; Francis and Hare 1994; Mann and Drinkwater
1994).
Over the last two decades, biological ‘‘output’’
variables have often covaried strongly and for
prolonged periods with variables that affect and
reflect the physical dynamics of the ocean. These
time series correlations suggest a close association
between marine populations and ‘‘ocean climate’’
(Mackas and de Young 2001). After Ebbesmeyer
et al. (1991) showed a consistent climate-related
abrupt shift around 1975–1979, based on the study of
40 environmental variables from the Pacific and the
Americas, another four regime shifts were identified
in 1890, 1925, 1947, and 1989 (Ebbesmeyer et al.
1991; Beamish et al. 1999; Watanabe and Nitta 1999;
Mantua 2001). Many climate indices were deter-
mined to have undergone a regime shift in 1977
contributing to ecosystem regime shifts world wide,
including SO (Southern Oscillation Index), PNA
(Pacific North American Index), ALPI (Aleutian Low
Pressure Index), NPI (North Pacific Index), Niño 3,
Niño 1 ? 2, Niño 3 ? 4, Niño 4, WP (west Pacific
Index), PDO (Pacific Decadal Oscillation), PCI
(Pacific Circulation Index), AO (Arctic Oscillation),
NAO (North Atlantic Oscillation), ACI (Atmospheric
Circulation Index for the Atlantic basin), and TNI
(Trans-Niño Index). At the same time, a number of
studies found synchronous changes in the environ-
ment and biological abundance. Catch and abundance
pattern in fish populations were used as a mirror of
climate-ocean regime shifts (Hare and Francis 1995;
Clark et al. 1999; Anderson 2000; McFarlane et al.
2000; Reid et al. 2001).
The potential implications for fisheries manage-
ment from the regional impacts of environmental
changes have been studied for many years. Though
overfishing was deemed to be responsible for the
shifts, there was historical evidence (Soutar and
Isaacs 1974; Cushing 1982; Francis and Hare 1994)
that supports the idea that these shifts occurred before
there was heavy fishing (Steele 1996). Evidence
therefore supports the assumption of regime shifts,
indicating that environmental factors are the
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Rev Fish Biol Fisheries (2009) 19:177–191
underlying mechanism, along with fishing effort
(Steele 1996).
Fish populations are influenced by many factors of
their natural environments during all phases of their
life cycles, but especially in their early life stages.
Subtle changes in key environmental variables, such
as temperature, salinity, wind, ocean currents, and
upwelling, can sharply alter the abundance, distribu-
tion, and availability of fish populations, either
directly or by affecting prey or predator populations.
Human activities can also affect the sustainability of
fish populations through the application of a variety
of different management schemes by using new
technologies, each of which could have a beneficial
or adverse consequence for the state of the fishery
(Glantz 1992). Such changes affect production
(especially recruitment) directly, and cause severe
confusion in management systems when assessment
scientists cannot distinguish between climatic and
harvesting impacts.
It was not until the ‘‘regime shift’’ concept was
used (Francis and Hare 1994; Hare and Francis 1995;
Mantua et al. 1997) that it became a popular topic in
ecosystem, fishery and climate-ecosystem research
(Steele 1996, 1998). PDOs (Pacific Decadal Oscilla-
tion) of 1977 and 1989 were regarded as regime shifts
through intervention and cross correlation analyses.
There were substantial changes during these two
periods in Climate Index and biological factors (fish
and plankton abundance, Fromentin and Planque
1996; Hare and Mantua 2000; Mantua and Hare
2002). However, David Pierce (Scripps Institute of
Oceanography) and Matthew Newman (NOAA–
CIRES Climate Diagnostics Center) drew the same
conclusion that the PDO during the last 50 years were
caused by slow variation in the chaotic evolution of
tropical ENSO episodes, with some amplification of
the decadal components by the long, slow responses
of the extratropical Pacific Ocean to the year-to-year
ENSO variations (Dettinger 2001). PDO is speculated
to originate in the tropics, and is not a climatic
oscillation independent from the extratropical North
Pacific (Pierce 2001; Newman et al. 2003). Decadal
variations of the Pacific climate are artifacts of slow
but essentially random variations of the global
climate. Pierce and Newman termed this ‘‘red-noise
climate’’, contrary to the concept of climate regime
shift (Pierce 2001; Newman et al. 2003).
Rev Fish Biol Fisheries (2009) 19:177–191
Marine ecosystem regime shift
Examples of regime shifts in ecosystems
In the northeast Pacific, large fluctuations of Califor-
nia sardine (Sardinops sagax caerulea) and anchovies
(Engraulis mordax) have been observed over the past
50 years, similar to their major recoveries and
subsequent collapses over the last 2000 years. The
current recovery is not unlike those of the past in
magnitude (Baumgartner et al. 1992). At the same
time, Pacific salmon from Alaska (Oncorhynchus
spp.) showed nearly synchronous changes across
different species as well as across a large portion of
their spatial range in 1976/1977 and the late 1940s/
early 1950s (Francis and Hare 1994; Mantua et al.
1997; Mantua and Hare 2002). Sablefish (Anoplop-
oma fimbria) exhibited decadal-scale patterns in the
relative success of year classes (King and McFarlane
2000). Pacific halibut (Hippoglossus stenolepis)
experienced large periods of increase and decrease
over the last 100 years (McCaughran 1996; Clark
et al. 1999). The great increase of halibut in 1977–
1978 was regarded as part of the same regime shift
related to climate change (Clark et al. 1999). Alaska
king crab (Paralithodes camtschatica), Tanner crab
(Chionoecetes bairdi), and pink shrimp (Pandalus
borealis), important crustaceans in the Gulf of
Alaska, have experienced severe changes in their
distributions and abundance during the past 45 years.
This could be due to climatic change in the ocean,
which may cause low recruitment or high natural
mortality (i.e., negative regimes) (Haaker et al. 1998;
Fu and Quinn 2000), overfishing, or a combination of
these factors. During periods of warming, the crus-
taceans decline, while finfish such as cod, pollock,
and flatfish increase in numbers. Zooplankton pro-
duction was found to have large and highly
significant interannual and interdecadal fluctuations
during summer in the subarctic North Pacific.
Between the late 1950s–early 1960s and the 1980s,
there were significant increases in zooplankton bio-
mass, regarded as characteristic of those climate-
driven regime shifts in ecosystems (Fromentin and
Planque 1996). Although there is no recent evidence
from phytoplankton in the Northeast Pacific, there is
evidence from Hawaiian waters (Haigh et al. 2001),
which showed a regime shift in 1976. The entire
biotic shift from plankton and shellfish to finfish
179
corresponded with the timing of the climatic shift
shown as abrupt changes of air temperature and
pressure, and water temperature and salinity.
Another important example of ecosystem regime
shift is in the northwest Atlantic Ocean. Northern cod
(Gadus morhua) experienced large fluctuations spa-
tially and temporally. Northern cod began to increase
sharply about 1960, but after a peak in 1968, landings
declined steadily for a decade, rose briefly through the
1980s, and recently plunged to the lowest level in this
century (Shelton et al. 1996). Other fishes, such as
haddock (Melanogrammus aeglefinus), red hake (Uro-
phychis chuss), and capelin (Mallotus villosus), and
invertebrates such as short-finned squid (Illex illece-
brosus), Atlantic lobster (Homarus americamus), and
sea scallops (Placopecten magellanicus), also experi-
enced large variations in landings (Koslow et al. 1987;
Rowell and Trites 1985; Williamson 1992; Hoffmann
and Powell 1998; Post et al. 1999), yet showed
synchronous changes over large geographic areas.
The southeast Australian ecosystem also experi-
enced impacts from climate changes, which were
somewhat different. Recent studies show that climate
is linked to changes in some of the fish stocks and
reefs in this area. Catches of Tasmanian rock lobster
(Jasus edwardsii), abalone species, and a variety of
finfishes have shown correlations with climate vari-
ation in local wind fields, sea surface temperatures
and surface productivity. Orange roughy (Hoploste-
thus atlanticus) showed substantial responses to
climate variability at 600–1,500 m depth (CVAP
1998). Reef bleaching is severe under El Niño
conditions. The time series data strongly suggest
two influences: variation in the strength and annual
persistence of regional winds; and a long-term
southward shift in the subtropical ridge, which is
assumed to affect rainfall, nutrient regeneration, and
primary productivity. It is unlikely to affect winds,
which have shifted poleward approximately 300 km
in the past 70 years (CVAP 1998).
Possible driving forces for marine ecosystem
regime shifts
Driving force 1: climate-ocean decadal oscillations
Climate changes has been regarded as one of the
major driving forces of marine ecosystem regime
shift (Benson and Trites 2002; Beamish et al. 2004;
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180
Lees et al. 2006). Some scientists prefer that the term
marine ecosystem regime shifts be strictly used when
linked to climatic changes (Beamish et al. 2004). The
major reason to link ecosystem regime shift to
climate changes is because of the consistent time
scales between ecosystem regime shifts and climate
regime shifts, as shown in many examples. As
discussed by Beamish and Mahnken (1999), regime
shifts are changes in state over 10–30 years, and
regimes are large, linked climate-ocean ecosystems.
Climate changes at decadal scales, from natural or
anthropogenic causes, are likely to produce or
enhance regime shifts.
Decadal regime shifts occur over spatial scales of
thousands of kilometers (Table 1, Fig. 1). They are
mainly influenced by physical processes of gyres
(e.g., Sargasso Sea), continental upwelling (e.g., Peru
Current), water mass boundaries (e.g., Antarctic
convergence), and climatic-oceanic oscillations
(e.g., ENSO, AO, NAO, and PDO independently or
in combination). Biological responses to the regime
shifts occur in regional communities and ecosystems.
Marine ecosystems are changeable, given such a
changeable environment as the ocean. Table 1 (from
Steele 1998) demonstrates that the similarity in time
scales between the physical and biological processes
in the ocean indicates that marine ecosystems are
closely coupled to the physical dynamics.
Figure 1 illustrates the major mechanisms that
induce changes in biota. Physical effects include
turbulence, vertical mixing, tidal mixing and run-off
on a scale of less than 1 km; fronts, upwelling and
tides, tidal mixing and internal waves on scales of
1–1,000 km; gyres, large currents, global climate
oscillations etc. on scales of thousands of meters; and
thermohaline circulation on scales of millions of
meters. Changes in ocean physics at decadal scales
can be a significant factor influencing regime shifts
Table 1 A schematic description gives the time scales of change for processes related to ecosystem variability (Steele 1998)
Process Time scale (years)
1 3
Biology Stock recruitment Stressed (overfished)
Physics El Niño prediction North Atlantic oscillation
Models Single species Multispecies
Management TAC Mesh size
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Rev Fish Biol Fisheries (2009) 19:177–191
on the shelf and in inshore waters (Hoffmann and
Powell 1998).
El Niño is a disruption of the ocean-atmosphere
system in the tropical Pacific having important
consequences for weather around the globe. During
El Niño, the trade winds relax in the central and
western Pacific leading to a depression of the
thermocline in the eastern Pacific, and an elevation
of the thermocline in the west. This reduces the
efficiency of upwelling to cool the surface and cuts
off the supply of nutrient rich thermocline water to
the euphotic zone in the eastern Pacific coast. The
result is a rise in sea surface temperature and a
drastic decline in primary production, adversely
affecting higher trophic levels of the marine food
chain, including commercial fisheries in the region.
In spite of the destructive nature of El Niño, some
marine creatures benefit from the disturbance, such
as purple snails (Plicopurpura pansa), octopuses
(Octopus sp.) and shrimp stocks of the eastern
Pacific coast (Forrester 1997; Arntz 1986). El Niño
and La Nina greatly impact the fisheries and
ecosystems of coastal Peru and Chile. The Austra-
lian and Indian Ocean, north Pacific and some other
extra tropic ecosystems are also greatly influenced
over decadal scales.
The PDO (Hare 1996; Mantua and Hare 2002)
has been described as a long-lived El Niño-like
pattern of Pacific climate variability. Major changes
in northeast Pacific marine ecosystems have been
correlated with phase changes in the PDO. Warm
eras enhanced coastal biological productivity along
the western North America shelf, while cold PDO
eras have the opposite impact on marine ecosystem
productivity. PDO influences the biomass, growth,
distribution, and migration of plankton, inverte-
brates, and fishes (Hare et al. 1999; Hare and
Mantua 2000).
10 30 100
/Regime shifts? Natural
Thermo-haline circulation
Community/ecosystem
Effort control
Rev Fish Biol Fisheries (2009) 19:177–191 181

Fig. 1 Schematic showing

the dominant space and
time scales in the ocean for
(a) physical motions and (b)
biological population. In (b)
the overlapping boxes to the
left represent typical size
ranges (on the x axis) and
typical times for population
doubling (on the y axis) for
each type of organism. The
boxes to the right present
typical space scales for each
organism during its lifetime.
Figure from Hoffmann and
Powell (1998)

NAO is a large-scale oscillation in atmospheric
conditions between the subtropical high located near
the Azores and the sub-polar low near Iceland.
Changes in the mass and pressure fields lead to
variability in the strength and pathway of storm
systems crossing the Atlantic from the US East coast
to Europe. The NAO is most noticeable during the
winter season (November–April) with maximum
amplitude and persistence in the Atlantic sector. A
positive NAO index results in warm and wet winters
in Europe and cold and dry winters in northern
Canada and Greenland, whereas negative NAO
conditions bring moist air into the Mediterranean
and cold air to northern Europe, and milder winters to
Greenland (Hurrell 1995). Communities and ecosys-
tems in north Atlantic are greatly influenced by it
(Drinkwater 2002).
The Arctic Oscillation has wide effects in the
Northern Hemisphere. Acceleration of a counter-
clockwise spinning ring of air around the polar region
could be responsible for warmer winters in Scandi-
navia and Siberia, thinning of the stratospheric ozone
layer, and significant changes in surface winds that
might have contributed to Arctic ice thinning (Strich-
erz 1999). Recent research shows that the NAO is
linked to the AO (Kerr 2000). The trend in the Arctic

123
182
Oscillation has been reproduced in climate models
involving simulated increased concentrations of
greenhouse gases (Fyfe et al. 1999).
Driving force 2: fishing
For the mechanisms of regime shifts in ecosystems,
most of the science focuses on climate-related
changes. There are also many other reasons that
may cause or maintain large marine ecosystem
changes. Fishing is regarded as another important
driving force of marine ecosystem regime shrifts
(Reid et al. 2000; Jackson et al. 2001; Daskalov
2002; Harvey et al. 2003).
Overfishing is especially severe on species of high
economic value, such as the top predators (Christen-
sen et al. 2003). A deteriorating environment
combined with overfishing may result in fishing
down below the level of minimum population
biomass. This kind of deterioration in a dynamic
ecosystem is hard to recover from, as indicated by
Pacific sardine around California, and Atlantic cod
around Newfoundland. Recent anomalies in the
periodicities of stock fluctuations were likely due to
intense fishing in an unfavorable climate regime
(Steele 1998; Hilborn et al. 2003; Lees et al. 2006).
Fishing pressure in the North Atlantic has been
regarded as the cause that obscures or distorts natural
multi-annual abundance swings (Bakun 2004).
Marine food webs have also changed in the last
half-century. From the trophic level trends in major
fishery areas (Pauly et al. 1998), a synchronous but
opposite trend was observed, which implied commu-
nity changes due to decadal regime shifts. At the
same time a common trend of trophic level decline
was observed, implicating worldwide overfishing.
Driving force 3: introduced species
Introduced species have long been recognized as a
source that can greatly change marine ecosystems
(Bakun 2004). Introduced species such as mollusks,
Spartina sea grasses and fishes have been observed to
cause large marine ecosystem changes (Thompson
1991). One famous example of great marine ecosys-
tem change resulting from an introduced species is
Potamocorbula amurensis in northern San Francisco
Bay. Potamocorbula amurensis is a benthic suspen-
sion-feeding bivalve native to Asian Pacific waters. It
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Rev Fish Biol Fisheries (2009) 19:177–191
was accidentally introduced in San Francisco Bay in
1986, and dispersed quickly throughout the estuary
with dramatic ecological consequences. Field and
laboratory evidence suggested that this species was
capable of consuming most of the phytoplankton
produced in northern San Francisco Bay (Colern and
Alpine 1991). Benthic and pelagic communities have
been altered. As with many changes due to intro-
duced species in freshwater, estuarine, and marine
ecosystems, it is hard for the ecosystem to reestablish
or recover.
Driving force 4: river flow
River flow has been a major driving force for coastal
ecosystem changes (Royer 1979; Mahe 1991; Chen
et al. 2000). In the Yellow Sea and Bohai Sea
drainages, terrestrial run-off has changed greatly in
the last 50 years because of climate and human-
induced reasons. The biological community in this
area has also changed greatly, and succession of
dominant species has been obvious among different
periods. Some coastal species, such as large icefish
(Protosalanx hyalocranius) and estuarine tapertail
anchovy (Coilia ectenes), which prefer low salinity
habitat, have disappeared (Chen et al. 2000). Nutrient
runoff has eutrophified the coastal waters, changing
the ecosystem greatly.
Driving force 5: other forces that can cause
or enforce large marine ecosystem changes
Other forces that are observed to cause or enhance
large marine ecosystem changes include pollution,
disease and eutrophication (Bakun 2004; Lees et al.
2006). The disease-driven collapse of the herbivorous
urchin Diadema antillarum is one important reason
that the Jamaica coral reef community converted
from a coral habitat to a seaweed habitat (Hughes
1994). Coastal eutrophication can cause extensive
hypoxia and anoxia, such as the 1987 summer of the
Long Island Sound, New York.
The complicated biological relationships among
fisheries and other aquatic biota and their physiolog-
ical responses to environmental change are not well
understood. As a result, clear understanding of their
impacts upon regime shifts is a problem. Multiple
regimes have been observed in some marine ecosys-
tems (Zhang et al. 2000; Overland et al. 2006; Steele
Rev Fish Biol Fisheries (2009) 19:177–191
2004). Regime shift patterns in different ecosystems
are still not well understood and hard to predict
through climatic-oceanic oscillation models (Zhang
et al. 2000; Pierce 2001).
Controversy or questions about the concept
of regime shift
Regime shift and red noise?
Regime shift is a concept of decadal or centennial
scale. In such a short time it is hard to determine
whether a shift has occurred or not over longer
periods. The term red noise and red-shifted noise
have been coined for noise with positive autocorre-
lation, which results in a dominance of low
frequencies in the spectral decomposition of the time
series (Rudnick and Davis 2003; Bakun 2004;
Overland et al. 2006). Generally, regimes occur on
decadal time scales (Beamish et al. 2000). Statistical
time series analyses have generally compared 20–
60 years of time series data, and regarded the shifts
among them as a regime shift. Although Francis and
Hare (1994) extracted white noise while analyzing
intervention, most of the works on climate fishery
analysis did not consider the chaos from red noise.
Many climatologists concluded that the recent cli-
mate and fishery changes are basically a random
fluctuation with red noise inside, that is, red-noise
climate and red noise ecosystem dynamics (e.g.,
Pierce 2001; Rudnick and Davis 2003). Detailed
statistical analyses on red noise in climate models are
provided by Sardeshmukh et al. (2001), Torrence and
Compo (1998), and Overland et al. (2006). An
alternative is to discard long-term time series anal-
yses, and only consider regime changes over decadal
time scales. Hare and Mantua (2000) introduced the
concept of abrupt changes at low frequency, which
helped to avoid the dispute about red noise.
How to separate impacts from different driving
forces in ecosystem regime shifts?
Although there is historical evidence of regime shifts,
recent periodicities of stock fluctuations have shown
differences in amplitude and cycle (e.g., North Pacific
sardine and anchovy, Peru anchovy, Atlantic cod) due
to heavy fishing. There are also other factors that
183
could induce ecosystem changes, including intro-
duced species, commercial fishing, and pollution.
These mechanisms have been recognized, and much
work has been done to prevent over-fishing and
pollution and to protect ecosystems (Bakun 2004;
Lees et al. 2006). While attempting to quantify these
impacts on ecosystems in assessment and manage-
ment, it is hard to differentiate among them (Reid
et al. 2000). More work is needed to enhance the
study of biological and physical processes in marine
ecosystems to best understand how to accurately
quantify them.
Marine ecosystem regime shift, good or bad?
Positive effects such as longer growing seasons,
lower natural winter mortality, and faster growth
rates of species in high latitude may be offset by
negative factors such as an abrupt changing climate
that alters established reproductive patterns, migra-
tion routes, and ecosystem relationships. Although
marine systems are much more responsive to decadal
scale alternations in their physical environment, they
are also much more adaptable to those alternations.
Regime shifts in fish communities can have major
economic consequences without being obvious eco-
logical disasters (Steele 1998): succession and
replacement are observed. There are obvious declines
in the landings of highly economic species, but there
are also increases in other fishes and bivalves, such as
the replacement of cod, haddock, and flounder by
dogfish, skate, and mackerel on Georges Bank off
New England (Fogarty and Murawski 1998), and the
replacement of small yellow croaker (Larimichthys
polyactis), large yellow croaker (Larimichthys croce-
a), and left eye flatfish (Paralichthys olivaceus) by
herring (Ilisha elongata) and noodlefish (Salangidae)
in the Yellow and Bohai Sea (Chen et al. 2000). The
new marine community is as diverse and ecologically
acceptable as the previous one, and in some places,
more diversified, though the economic value may be
lower.
Some natural or human induced factors cause
switches in fish communities over time scales of a few
decades. Coastal domain areas are easily impacted by
human activities through addition of nutrients, drain-
age, land development, and by fish and shellfish
farming. ‘‘Restoration’’ of a marine ecosystem is
basically impossible (Steele 1998), as it is difficult to
123
184
determine the pristine, pre-disturbance state. Conti-
nental shelves are the regions where approximately
85% of fish landings originate, and where most
questions arise, including environmental changes on
fisheries and fishing impacts on marine environments.
In the deep ocean there is large interannual variability
in physical processes with strong indications of
longer-term trends in both biomass and species
composition. Changes in major current systems alter
the basic productivity and also have significant
consequences for marine communities in the ocean
(Mann and Lazier 1996).
Models in simulating and predicting ecosystem
regime shift
Because of the wide influence of regime shift in
ecosystems, modeling this phenomenon becomes
important in marine ecosystem study. Our ability to
manage marine ecosystems will increase with better
modeling quality. There are two basic kinds of models
in simulation and predicting regime shifts: historical
descriptive models and general circulation models
(GCMs). The first type uses analogies, descriptively
or statistically approaching the issue of scale through
the interpretation of patterns in time and space. The
second method simulates the general circulations in
climate-ocean or climate-ocean-biotic systems to
describe the climate and ecosystem dynamics.
Time series methods of intervention analysis
(Francis and Hare 1994; Hare and Francis 1995;
Mantua et al. 1997; Zhang et al. 1997), cross corre-
lation analysis (Francis and Hare 1994), Auto
Regressive Integrated Moving Average (ARIMA)
(Francis and Hare 1994; Hare and Francis 1995;
Mantua et al. 1997; Zhang et al. 1997), Cumulative
Summation (CuSum) (Murdoch 1979; Ebbesmeyer
et al. 1991; Hare and Mantua 2000) and red noise
models (Pierce 2001; Rudnick and Davis 2003;
Overland et al. 2006) have been used in the research
of historical descriptive models. Correlation associ-
ations have a disturbing tendency to break down part
way through long time series (Mackas and de Young
2001).
Atmospheric GCMs have been used to address how
the regional climate associated with monsoon rain-
fall and circulation anomalies responds to imposed
SST anomalies. Difficulties exist: for example,
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Rev Fish Biol Fisheries (2009) 19:177–191
17 atmospheric modeling groups in the Monsoon
Numerical Experimentation Group (MONEG) of the
world Climate Research Programme ran a 90-day
integration using the observed global SST fields for
1987 and 1988 as boundary forcing and atmospheric
fields as initial conditions. The results were not ideal
and suggested that the Indian Ocean region is difficult
to model. The physical mechanisms that caused the
PDO are not currently known (Mantua and Hare
2002). Despite the difficulties of developing GCMs,
several studies have appeared in recent years (Werner
et al. 2001; de Young et al. 2004).
No matter if one adheres to the regime shift
concept that there are relatively rapid changes in
community structure, or accepts only the probability
of auto-correlation around decadal time scales, it is
evident that predictions based on uncorrelated, white
noise, recruitment will degrade rapidly. Presently,
with a short history of ecosystem and climate data, it
is effective to regard this kind of change in ecosys-
tems as a process whereby the ecosystem changes
from one stable state to another stable state because
of human-induced or natural causes. Considering new
types of recruitment and growth patterns in making
assessments is necessary where old patterns fail
(Steele 1996).
It is important to note that forecasting by analogy
is not an attempt to assess the direct effects of a
climate change on the biological aspects of living
marine resources: correlation does not imply causal-
ity. The physical-biological mechanisms that drive
the observed correlations need to be understood in
order to identify a stable predictive scheme and
therefore be effective in a time of changing back-
ground climate.
Recent works have sought the key mechanism or
link between climate and oceanic ecosystem output.
Polovina et al. (1995) concluded from their research
that changes in mixed layer depths occurred on
decadal and basin scales and could be a key
mechanism linking variation in the atmosphere and
oceanic ecosystems. Climate warming was regarded
as the key mechanism that caused the decline of
zooplankton in the California Current (Roemmich
and McGowan 1995). Higher surface temperature
increased the thickness of the discontinuity layer,
which resulted in less lifting of the thermocline by
wind-driven upwelling, and subsequently reduced the
inorganic nutrient supply for zooplankton. It is
Rev Fish Biol Fisheries (2009) 19:177–191
therefore important to explore mechanisms in specific
regional marine ecosystems in order to specify the
specific mechanism in that region.
The regime shift concept also implies that different
regimes have an inherent stability, so that significant
forcing is required to flip the system to an alternative
state (Steele and Henderson 1984). Regime shift
models usually have only two states, which are
usually not the real systems because environmental
changes or human forcing may result in unintended
consequences. Many kinds of fish have disappeared
from marine ecosystems, and may never come back,
and lagged effects have appeared for some species.
As a result, some ecosystems have shown more than
two regimes during dynamic time series (Zhang et al.
2000; Bakun 2004).
The production of fish biomass in the oceans is
governed by interactions among numerous physical,
chemical, and biological processes. It is hard to
define an effective GCM to be used for climate-
related impact analyses. The key physical mecha-
nisms of the ocean that impact biological factors are
not sufficiently studied. Many ocean systems or
domains can provide more production, such as
upwelling systems for pelagic fishes, phytoplankton,
and zooplankton, downwelling domains for bottom
fishes, mixing of runoff and ocean sea water, mixing
of warm and cold currents, and advection (Ware and
McFarlane 1989; Ware and Thomson 1991; Roem-
mich and McGowan 1995; Wickett 1967). Analyses
are qualitative and not included in the ecosystem
models and fishery assessment.
At present, the spatial resolution of GCMs is too
coarse to be useful for reliable and credible regional
impact assessment. When computers become more
powerful and our understanding of the physical
process increases, we will rely more and more on
the dynamic forecasts of GCMs. They have the
tremendous advantage of working forward from
the actual present observed conditions, thus avoiding
the problem of statistically averaging over a number
of events that differ in important details.
Models in simulating and predicting regime shifts
may fail because of co-impacts from natural and
human-induced reasons. Rigler (1982) concluded that
an inability of empirical science to provide reliable
predictions or forecasts of future states of both
ecological and connected weather systems has led to
a general resistance to historical science. He also said
185
that this is not the fault of the method: rather, it is
because ‘‘long-term abundance of species in systems
subject to anthropogenic or other changes is not
predictable.’’
From Table 1 (Steele 1998), we can see that
annual and decadal perspectives are not alternatives,
but are complementary. Assimilating different types
of research (single species, multispecies, and ecosys-
tem) and assessment approaches into a single
scientific program or management system is impor-
tant for future work. This helps to improve
understanding, assessment and management of our
marine resources under the situation of regime shifts.
Impact of regime shifts on fisheries and difficulties
in integrating regime shifts into fisheries science
Climate changes affect marine ecology through
alteration of nutrient cycles, primary production, fish
growth, recruitment, and community changes. Fish-
eries certainly are affected, as recruitment (Mann and
Drinkwater 1994; Wada and Jacobson 1998), and
growth (DeAngelis et al. 1991; Hutchings 1999;
Clark et al. 1999) are heavily impacted. Changes in
catches, dominant species, composition, replacement,
and distribution are widely observed in world fisher-
ies (Mann and Drinkwater 1994; Arntz 1986;
Beamish et al. 1999). Use of climate information in
fisheries and ecosystem models is increasing to
identify methodological and theoretical gaps in
current knowledge and explore the possibilities for
resolving them. This information is also being used to
qualify the ecological impact with a view toward
incorporating climate information within operational
decision making in fishery management. In addition,
global collapses of fish stocks were likely affected not
only by ocean climate changes and fishing, but were
also influenced by many other factors, such as
pollution (Wolfe 1985), introduced species, and other
human activities.
Most global commercial fisheries have been
affected by decadal climate changes. Commercial
fishing catches can provide a measure of the ecosys-
tem changes. The causes of collapses of the California
sardine and anchovy have been argued for decades,
including overfishing, environmental fluctuation, or
competition for food resources between sardine and
anchovies. Analysis of fossil sardine time series has
123
186
revealed that sardine production has fluctuated natu-
rally (Soutar and Isaacs 1974); large, natural, pre-
historical fluctuations were clearly unrelated to com-
mercial fishing. Spectral analysis of the scale-
deposition series off southern California from A.D.
270 through 1970 showed that sardines and anchovies
both tend to vary over a period of approximately
60 years. It also revealed nine major recoveries and
subsequent collapses of the sardine population over
1,700 years. The current recovery is not unlike those
of the past in magnitude (Baumgartner et al. 1992).
Through intervention analysis, the climate index of
Kodiak Winter Air Temperature and North Pacific
Index, and Alaska salmonid production showed
nearly synchronous interventions across different
species, as well as across a large part of the spatial
range in 1979 and late 1940s/early 1950s. Mantua
et al. (1997) analyzed climate indices in the North
Pacific Ocean along with regional climate indices
within salmonid habitats. Their work showed inter-
vention in the time series data and strong correlation
among them.
Peruvian anchovy and Chilean sardine was one of
the earliest fisheries realized by fishermen and
scientists to have large fluctuations. This fishery is
one of the most productive in the world. During the
1960s and early 1970s, catches in this area comprised
nearly 20% of the world’s landings. Catch data are
available from 1950 to the present. This time series
showed that catch stayed low until 1958, then sharply
increased in 1958–1962, when the annual catch
reached 7,000,000 tonnes. Subsequently, it grew to
12,000,000 tonnes in 1970, suffered a major collapse
in 1972, and then showed a slow and uneven recovery
process. The 1972 collapse was considered to have
been prompted by an El Niño event. There was also a
synchronous change between fish catch and climatic
conditions, which leads to the prospect of probabi-
listic catch forecasts of anchovy constructed on the
basis of ENSO forecasts. However, climate variabil-
ity is not a factor that can be linearly translated into
absolute catch values. Historical, political, and
perhaps circumstantial factors also play roles (Glantz
1992; Hilborn et al. 2003).
Another important point concerning the impact
from regime shifts is that most of the time we only
pay attention to the negative impact, but responses of
biological factors to climate changes are not always
negative. Even in El Niño periods, when anchovy was
123
Rev Fish Biol Fisheries (2009) 19:177–191
reduced, sardine and jack mackerel increased (Glantz
1992). In the northeast Pacific marine ecosystem,
warm PDO phases have favored high salmon pro-
duction in Alaska and low salmon production off the
west coast of California, Oregon, and Washington.
Conversely, cool PDO eras have favored low salmon
production in Alaska and relatively high salmon
production for California, Oregon, and Washington
(Hare 1996; Hare et al. 1999). The community/
ecosystem changes when the climate changes, but is
still functional though the economic influence may be
heavy (Steele 1996).
The goal of fishery management is to predict next
year’s fish biomass through models, which may
include backward Virtual Population Analysis
(VPA), and forward recruitment, growth, and mor-
tality predictions. For management purposes,
production theory, yield-per-recruitment and spaw-
ner-recruitment models are widely used to conduct
best policy searches (Quinn and Deriso 1999). No
matter which kind of model is used, historical data
are needed. After a population dynamic model is
established, data are fitted to the model, different
management options are tested to predict future
biomass and to get the best management limit. The
classical population dynamic models usually assume
that the environment is an averaged constant, because
it is hard to account for dynamic patterns and
intervening variables; these issues have been ignored
in stock assessment using either VPA analysis or
forward abundance prediction. Production models
ignore year to year changes in abundance. Finally,
yield per recruitment models unrealistically assume
that recruitment is constant from year to year
(Hilborn and Walters 1992; Quinn and Deriso 1999).
Climate can have substantial impacts on various
stages in the life cycle of many species, and can alter
primary production, triggering changes that propa-
gate through the trophic web. Relevant impacts can
occur from very local to basin-wide spatial scales,
and on timescales ranging from hourly to interdec-
adal. Recruitment changes triggered by
environmental changes are widely recognized, such
as the recruitment of Atlantic cod fisheries after 1989.
At the same time changes in growth, natural mortal-
ity, distribution and immigration are also observed in
many fisheries (Drinkwater 2002).
Regime shifts are therefore quite contrary to the
common assumption of stable state, around which
Rev Fish Biol Fisheries (2009) 19:177–191
there lies a normal distribution of variation, in
fisheries science. Conventional statistical analyses,
which assume linearity and normality in a set of
observations, are inadequate in understanding under-
lying mechanisms that link climate, ocean and biotic
systems across possible regimes. It is therefore
necessary to have comprehensive knowledge of both
short-term and long-term effects, including climate
changes, regime shifts, changes in fish stocks and
feedback of biological systems to physical events
(Beamish et al. 1999).
As for ecosystem models discussed in the section
of modeling marine ecosystem regime shrift, there are
also two approaches for incorporating environmental
data into stock assessment. One is to add environ-
mental parameters into classical stock assessment
models directly (e.g., Jacobson and MacCall 1995).
Another is to build biological-physical process mod-
els (Bakun 1996; Haigh et al. 2001; Wiebe et al.
2001). The first approach is practical, based on the
present knowledge and data available. However, there
are problems: correlation does not mean causality;
correlation associations are not always consistent
along time series; and climate variability cannot be
linearly translated into absolute catch value because
of this non-consistency. The second approach
involves marine ecosystem models. It is hard to
define an effective GCM to be used for climate-related
impact analyses, and it is hard to separate the impacts
from fishing and climate changes. At the same time, in
the present situation of low-resolution climate data,
GCMs are insufficient for assessment of fish popula-
tion processes at regional scales.
Research needs to incorporate regime shifts in
modeling fish dynamics, although there are some
studies that incorporate environmental factors into
classical stock assessment directly, or use GCMs to
predict population biomass (Werner et al. 1996,
1997, 2001). As it is difficult to find key environ-
mental factors attributed to the regime shift, an option
is to use classical models but different parameters to
simulate different regimes. This requires accurate
forecasting of decadal climate change. Until GCM or
a well performing correlation model can be devel-
oped, an alternative is to discard long term time series
statistical analysis and only consider regime shifts
over decadal time scales. By doing this the concept of
ecosystem regime shift is useful from a management
standpoint even if the climate regime shifts cannot be
187
accurately predicted (Beamish et al. 2004). Precau-
tionary reference points with a risk assessment that
fully consider uncertainty of the ecosystem variabil-
ity should improve future management (Jiao et al.
2005).
Potential future research
More work is needed to improve our understanding of
regime shift and its impacts on ecosystems and
implications for fishery management. At the same
time it is important to reconsider the regime shifts
from macroscopic and microcosmic viewpoints. The
essence of ecosystem sensitivity and adaptability
requires extension of research on biological-physical
processes in regional fisheries.
The terms stability, sustainability, and restoration
are widely used in ecosystem and fisheries research.
The concept of the marine protected area is also a
popular term in ecology and fisheries (Pauly et al.
2002). In some situations, it can be helpful in keeping/
increasing biodiversity and/or productivity to provide
disturbances to the ecosystem (Lessard et al. 2005).
From an economic viewpoint, it is also practical to
induce disturbance to an ecosystem to get the
maximum benefit without destroying the ecosystem.
Fish landings are correlated with climate indices
(SST, inflow, salinity, pressure), but are also corre-
lated with predator abundance, water current, and
biomass of phytoplankton and zooplankton. Which
is the key mechanism: insufficient plankton as
food, Ekman influences on larval fishes, changed
temperature and salinity, other physical/chemical
circumstances, or co-impacts? Finding the answers
should be of high priority in current and future
research as classical population assessment models
are updated to incorporate climate impact and GCMs.
Traditionally, we have focused on yearly recruit-
ment to individual fish stocks. The possibility of
fishing down marine food webs requires comparison
of changes in community structure over time scales
of a few years in highly over-fished regions to longer
historical scales in future work (Steele 1998). It is
time to consider questions from an ecosystem view-
point. Present management failures call for deeper
understanding of fishing effects in the long term, such
as the ability of stocks to survive a number of years of
adverse environmental conditions.
123
188
Recent research has related the fisheries phenom-
ena of strong and weak recruitment/productivity to
the noise in nature. Red noise has been used in the
explanation of regime shift (Pierce 2001). The
classification of noise by spectral density is given
‘‘color’’ terminology, extending the concept of red
noise. White noise means a constant spectral density
and temporally uncorrelated noise signal. For colored
noise, the spectral density changes with changing
frequency, and the noise signal is auto-correlated
(Halley 1996; Petchey 2000). Noises in nature may
not be white, but colored in many cases (Halley 1996;
Caswell and Cohen 1995; Vasseur and Yodzis 2004).
Colored noises have more serious implications for
endangered species and species with low population
size (Halley and Kunin 1999; Morales 1999; Schwa-
ger et al. 2006). Before we are able to simulate the
processes using GCMs, classical stock assessment
model with colored noises may lead to better
inferences about the population size and the fishery
status (Schwager et al. 2006; Jiao et al. 2008).
More work is required on research techniques,
such as climate forecast systems. Detailed and high-
resolution data are needed for both GCMs and
classical stock assessment. Satellite information is
an effective tool in climate forecasting and biolog-
ical-physical analysis, which provides far superior
spatial coverage and in situ information. Forecasts of
El Niño clearly have direct value for fisheries in the
productive regions of western South America. More
accurate prediction of El Niño will also be very
valuable for countries outside of the tropics, such as
Japan, Canada and the United States. It can also help
fisheries managers to distinguish between changes in
populations due to anthropogenic factors and changes
from natural conditions. The application of satellite
information will be widely needed in both climate
and ecosystem research. Understanding the linkages
between physical environmental and biological fac-
tors requires input from a variety of disciplines,
coordinated research programs, and considerable
cooperation at national and international levels.
The works previously done on ecosystem regime
shift are at ocean basin scales, e.g., Mantua et al.
(1997) and Zhang et al. (2000). These large changes
in ecosystems are world-wide, as indicated by the
catch trajectories of some sympatric world fisheries.
Catch trajectories of many fisheries from west and
east Pacific, Atlantic and Australia waters show
123
Rev Fish Biol Fisheries (2009) 19:177–191
regime patterns. At the same time, given the contro-
versy over independence between PDO and NAO, it
seems that climate oscillation may be world-wide.
PDO may be the result of joint action on the North
Pacific by El Niño and AO. NAO may express AO in
the North Atlantic. World ecosystems are all influ-
enced by oscillations of El Niño, AO, and Antarctic
Oscillation. Every basin scale and ocean scale
ecosystem, because of their distinctive geographic
characteristics, may show delayed significant to
inconspicuous ecological changes at similar or
different time scales. From this point of view, it is
time to consider climate GCMs as world climatic-
oceanic oscillations, not limited to basin and ocean
scales. Ecosystem models will benefit from accurate
climate oscillation predictions, but at present, con-
sideration of the key mechanisms that influence the
regional ecosystems is required.
Conclusion
Climate-ocean impacts have traditionally been
ignored in stock assessment, but are now known to
be important. Stock assessments and stewardship of
fisheries now require a distinction between fishing
effects and climate-driven effects. For now it is not an
issue of which approach is most appropriate, but
rather that there is a requirement to both provide
annual assessments and improve understanding of
natural processes. More effective strategies for fish-
eries will be provided. With the development of
science and technology, the mechanisms of climate-
driven effects will be clearer and GCMs will be more
accurate (de Young et al. 2004).
Acknowledgements I would like to thank Dr. Yong Chen of
the University of Maine, Dr. Joe Wroblewski, Dr. David
Schneider and Dr. Paul Snelgrove of Memorial University of
Newfoundland who provide valuable advice. This research was
supported by grants from Virginia Polytechnic Institute and
State University, and the USDA Cooperative State Research,
Education and Extension Service, Hatch Project #0210510, to
Y. Jiao.
References
Anderson PJ (2000) Pandalid shrimp as indicators of ecosys-
tem regime shift. J Northwest Atl Fish Sci 27:1–10. doi:
10.2960/J.v27.a1
Rev Fish Biol Fisheries (2009) 19:177–191
Arntz WE (1986) The two faces of El Niño 1982–1983. Me-
eresforschung 31:1–46
Bakun A (1996) Patterns in the ocean. California Sea Grant,
CIB, San Diego
Bakun A (2004) Regime shifts. In: Robinson AR, Brink K
(eds) The sea, vol 13. Harvard University Press, Cam-
bridge, pp 971–1018
Baumgartner TR, Soutar A, Ferreira-Bartrina V (1992)
Reconstruction of the history of Pacific sardine and
northern anchovy population over the past two millennia
from sediments of the Santa Barbara Basin, California.
CCOFI Rep 33:24–40
Beamish RJ, Mahnken C (1999) Taking the next step in fish-
eries management. Ecosystem approaches for fisheries
management. Alaska Sea Grant College Program, AK-
SG-99-01, pp 1–21
Beamish RJ, Noakes DJ, McFarlane GA et al (1999) The
regime concept and natural trends in the production of
Pacific salmon. Can J Fish Aquat Sci 56:516–526. doi:
10.1139/cjfas-56-3-516
Beamish RJ, McFarlane GA, King JR (2000) Fisheries clima-
tology: understanding decadal scale processes that
naturally regulate British Columbia fish population. In:
Harrison PJ, Parson TR (eds) Fisheries oceanography: an
integrative approach to fisheries ecology and manage-
ment. Blackwell Science, Oxford, pp 94–139
Beamish RJ, Benson AJ, Sweeting RM et al (2004) Regimes
and the history of the major fisheries off Canada’s west
coast. Prog Oceanogr 60:355–385. doi:10.1016/j.pocean.
2004.02.009
Benson AJ, Trites AW (2002) Ecological effects of regime
shifts in the Bering Sea and eastern North Pacific. Fish
Fish 3:95–113. doi:10.1046/j.1467-2979.2002.00078.x
Caswell H, Cohen JE (1995) Red, white and blue: environmental
variance spectra and coexistence in metapopulations.
J Theo Biol 176:301–316
Chen DG, Jiao Y, Liu Q et al (2000) Fish species diversity in
the Yellow River estuary. Acta Oceanol Sin 19:125–135
Christensen V, Guénette S, Heymans JJ et al (2003) Hundred-
year decline of North Atlantic predatory fishes. Fish Fish
4:1–24. doi:10.1046/j.1467-2979.2003.00103.x
Clark WG, Hare SR, Parma AM et al (1999) Decadal changes
in growth and recruitment of Pacific halibut (Hippoglos-
sus stenolepis). Can J Fish Aquat Sci 56:242–252. doi:
10.1139/cjfas-56-2-242
Colern J, Alpine (1991) Potamocorbula amurensis, a recently
introduced Asian clam, has had dramatic effects on the
phytolankton biomass and production in northern San
Francisco Bay. J Shellfish Res 10:258–259
Cushing DH (1982) Climate and fisheries. Academic Press,
New York
CVAP (Climate Variability in Agriculture R&D Program)
(1998) Climate and fisheries on the south-east Auatralian
continental shelf and slope. http://www.cvap.gov.au.new
fskoslow.html
Daskalov G (2002) Overfishing drives a trophic cascade in the
Black Sea. Mar Ecol Prog Ser 225:53–63. doi:10.3354/
meps225053
de Young B, Heath M, Werner F et al (2004) Challenges of
modeling ocean basin ecosystems. Science 304:1463–
1466. doi:10.1126/science.1094858
189
DeAngelis DL, Godbout L, Shuter BJ (1991) An individual-
based approach to predicting density-dependent dynamics
in smallmouth bass populations. Ecol Modell 57:91–115.
doi:10.1016/0304-3800(91)90056-7
Dettinger M (2001) Drought, epic droughts & droughty cen-
turies—lesson from California Paleoclimatic record. IEP
Newsl 14:51–54
Drinkwater KF (2002) A review of the role of climate vari-
ability in the decline of northern cod. Fish Soc Symp
32:113–130
Ebbesmeyer CC, Cayan DR, McClain DR et al. (1991) 1976
Step in Pacific climate: forty environmental changes
between 1968–1975 and 1977–1984. In: Betancourt JL,
Tharp VL (eds) In: Proceedings of the 7th annual Pacific
climate (PACLIM) workshop. Interagency Ecological
Study Program Technical Report 26, California Depart-
ment of Water Resources, pp 115–126, April 1990
Fogarty NJ, Murawski SA (1998) Large-scale disturbance and
the structure of marine systems: fisheries impacts on
Georges Bank. Ecol Appl 8:S6–S22
Forrester AL (1997) The effects of El Niño on marine life.
http://www.csa.com/hottopics/elNiño/overview.html
Francis RC, Hare SR (1994) Decadal-scale regime shifts in the
large marine ecosystems of the North-east Pacific: a case
for historical science. Fish Oceanogr 3:279–291. doi:
10.1111/j.1365-2419.1994.tb00105.x
Fromentin JM, Planque B (1996) Calanus and environment in
the eastern North Atlantic 2. Influence of the North
Atlantic oscillation on C. finmarchicus and C. helgo-
landicus. Mar Ecol Prog Ser 134:111–118. doi:
10.3354/meps134111
Fu C, Quinn TJII (2000) Estimability of natural mortality and
other population parameters in a length-based model:
Pandalus borealis in Kachemak Bay, Alaska. Can J Fish
Aquat Sci 57:2420–2432. doi:10.1139/cjfas-57-12-2420
Fyfe JC, Boer GJ, Flato GM (1999) The Arctic and Antarctic
oscillations and their projected changes under global
warming. Geophys Res Lett 26:1601–1604. doi:10.1029/
1999GL900317
Glantz MH (1992) Climate variability, climate change and
fisheries. Cambridge University Press, Cambridge
Haaker PL, Parker DO, Barsky KC et al (1998) Growth of red
abalone, Haliotis rufescens (Swainson), at Johnsons Lee,
Santa Rosa Island, California. J Shellfish Res 17:747–753
Haigh SP, Dennis DG, Hsieh WW (2001) Simulation of the
planktonic ecosystem response to pre- and post-1976
forcing in an isopycnic model of the North Pacific. Can J
Fish Aquat Sci 58:703–722. doi:10.1139/cjfas-58-4-703
Halley JM (1996) Ecology, evolution and 1/f-noise. Trends
Eco Evol 11:33–37
Halley JM, Kunin WE (1999) Extinction risk and the 1/f family
of noise models. Theo Pop Biol 56:215–230
Hare SR (1996) Low frequency climate variability and salmon
production. Ph.D. dissertation. School of Fisheries, Uni-
versity of Washington, Seattle
Hare SR, Francis RC (1995) Climate change and salmon pro-
duction in the northeast Pacific Ocean. In: Beamish RJ
(ed) Climate change and northern fish populations. Can J
Fish Aquat Sci (Spec Publ) 121: 357–372
Hare SR, Mantua NJ (2000) Empirical evidence for North
Pacific climatic regime shifts in 1977 and 1989. Prog
123
190
Oceanogr 47:103–145. doi:10.1016/S0079-6611(00)00
033-1
Hare SR, Mantua NJ, Francis RC (1999) Inverse production
regimes: Alaskan and west coast salmon. Fisheries 24:6–14.
doi :10.1577/1548-8446(1999)024\0006:IPR[2.0.CO;2
Harvey CJ, Cox SP, Essington TE et al (2003) An ecosystem
model of food web and fisheries interactions in the Baltic
Sea. ICES J Mar Sci 60:939–950. doi:10.1016/S1054-
3139(03)00098-5
Hilborn R, Walters C (1992) Quantitative fisheries stock
assessment: choice, dynamics, and uncertainty. Chapman
and Hall, New York
Hilborn R, Branch TA, Ernst B et al (2003) State of the world
fisheries. Annu Rev Environ Resour 28:359–399. doi:
10.1146/annurev.energy.28.050302.105509
Hoffmann EE, Powell TM (1998) Environmental variability
effects on marine fisheries: four case histories. Ecol Appl
8(Suppl):S23–S32
Hughes TP (1994) Catastrophes, phases shifts and large scale
degradation of a Caribbean coral reef. Science 16:
149–177
Hurrell JW (1995) Decadal trends in the North Atlantic
Oscillation: regional temperatures and precipitation. Sci-
ence 269:676–679. doi:10.1126/science.269.5224.676
Hutchings JA (1999) Influence of growth and survival costs of
reproduction on Atlantic cod, Gadus morhua, population
growth rate. Can J Fish Aquat Sci 56:1612–1623. doi:
10.1139/cjfas-56-9-1612
Isaacs JD (1976) Some ideas and frustrations about fishery
science. Calif Coop Ocean Fish Invest Rep 18:34–43
Jackson JBC, Kirby MX, Berger WH et al (2001) Historical
overfishing and the recent collapse of coastal ecosystems.
Science 293:629–637. doi:10.1126/science.1059199
Jacobson LD, MacCall AD (1995) Stock-recruitment models
for pacific sardine (Sardinops sagax). Can J Fish Aquat
Sci 52:566–577. doi:10.1139/f95-057
Jiao Y, Chen Y, Wroblewski J (2005) An application of
composite risk assessment method in assessing fisheries
stock status. Fish Res 72:173–183. doi:10.1016/j.fishres.
2004.11.003
Jiao Y, Neves R, Jones J (2008) Models and model selection
uncertainty in estimating growth rates of endangered
freshwater mussel populations. Can J Fish Aquat Sci 65
(in press)
Kerr R (2000) A new force in high latitude climate. Science
284:241–242. doi:10.1126/science.284.5412.241
King JR, McFarlane (2000) Decadal scale patterns in the rel-
ative year class success of sablefish, Anoplopoma fimbria.
Fish Oceanogr 9:62–70. doi:10.1046/j.1365-2419.2000.
00122.x
Koslow JA, Thompson KR, Silvert W (1987) Recruitment to
northwest Atlantic cod (Gadus morhua) and Haddock
(Melanogrammus aeglefinus) stocks: influence of stock
size and climate. Can J Fish Aquat Sci 44:26–39. doi:
10.1139/f87-004
Lees K, Pitois S, Scott C et al (2006) Characterizing regime
shifts in the marine environment. Fish Fish 7:104–127
Lessard RB, Martell S, Walters CJ et al (2005) Should eco-
system management involve active control of species
abundances? Ecol Soc 10:1. http://www.ecologyand
society.org/vol10/iss2/art1
123
Rev Fish Biol Fisheries (2009) 19:177–191
Mackas DL, de Young B (2001) GLOBEC Canada 1996–2000:
A sampler. Can J Fish Aquat Sci 58:645–646. doi:
10.1139/cjfas-58-4-645
Mahe G (1991) La Variabilite des apports fluviaux au golfe de
Guinea utilisee comme indice climatique. In: Cury P, Roy
C (eds) Pecheries Ouest-Africaines Variabilite, Instabilite
et Changement, pp 343–353
Mann KH, Drinkwater KF (1994) Environmental influences on
fish and shellfish production in Northwest Atlantic.
Environ Rev 2:16–32
Mann KH, Lazier JRN (1996) Dynamics of marine ecosystems:
biological-physical interactions in the oceans. Blackwell
Science, Oxford, p 394
Mantua NJ (2001) The Pacific decadal oscillation. In: McC-
racken MC, Perry JS (eds) The encyclopedia of global
environmental change, V1. The earth system: physical and
chemical dimension of global environmental change. pp
592–594
Mantua NJ, Hare SR (2002) The Pacific decadal oscillation. J
Oceanogr 58:35–44. doi:10.1023/A:1015820616384
Mantua NJ, Hare SR, Zhang Y et al (1997) A Pacific interdecadal
climate oscillation with impacts on salmon production. Bull
Am Meteorol Soc 78:1069–1080. doi :10.1175/1520-
0477(1997)078\1069:APICOW[2.0.CO;2
McCaughran DA (1996) Seventy-five years of halibut man-
agement success. In: Hancock DA, Smith DC, Grant A,
Beumer JP (eds) Second world fishery congress. CSIRO,
Brisbane, Australia, pp 680–686
McFarlane GA, King JR, Beamish RJ (2000) Have there been
recent changes in climate? Ask the fish. Prog Oceanogr
47:147–169. doi:10.1016/S0079-6611(00)00034-3
Merriam-Webster’s Collegiate Dictionary http://www.m-w.
com/cgi-bin/dictionary
Morales JM (1999) Viability in a pink environment: why
‘‘white noise’’ models can be dangerous. Eco Lett 2:
228–232
Murdoch J (1979) Control charts. MacMillian, London
Newman M, Compo GP, Alexander MA (2003) ENSO-forced
variability of the Pacific decadal oscillation. J Clim
16:3853–3857. doi :10.1175/1520-0442(2003)016\3853:
EVOTPD[2.0.CO;2
Overland JE, Percival DB, Mofjeld HO (2006) Red noise and
regime shifts in the North Pacific. Deep Sea Res Part I
Oceanogr Res Pap 53:582–588. doi:10.1016/j.dsr.2005.
12.011
Pauly D, Christensen J, Dalsgaard J et al (1998) Fishing down
marine food webs. Science 279:860–863. doi:10.1126/
science.279.5352.860
Pauly D, Christensen V, Guenette S et al (2002) Towards
sustainability in world fisheries. Nature 418:689–695. doi:
10.1038/nature01017
Petchey OL (2000) Environmental colour affects aspects of
single-species population dynamics. Proc R Soc Lond B
267:747–754
Pierce DW (2001) Distinguishing coupled ocean-atmosphere
interactions from background noise in the North Pacific.
Prog Oceanogr 49:331–352. doi:10.1016/S0079-6611
(01)00029-5
Polovina JJ, Mitchum GT, Evans GT (1995) Decadal and
basin-scale variation in mixed layer depth and the impact
on biological production in the Central and North Pacific,
Rev Fish Biol Fisheries (2009) 19:177–191
1960–1988. Deep Sea Res 42:1701–1716. doi:10.1016/
0967-0637(95)00075-H
Post E, Forchhammer MC, Stenseth NC (1999) Population
ecology and climate change: consequences of the North
Atlantic Oscillation (NAO). Econ Bull 47:117–125
Quinn TJ, Deriso RB (1999) Quantitative fish dynamics.
Oxford University Press, New York
Reid PC, Battle EJV, Batten SD, Brander KM (2000) Impact of
fisheries on plankton community structure. ICES J Mar
Sci 57:495–502. doi:10.1006/jmsc.2000.0740
Reid PC, de Fatima Borges M, Svendsen E (2001) A regime
shift in the North Sea circa 1988 linked to changes in the
North Sea horse mackerel fishery. Fish Res 50:163–171.
doi:10.1016/S0165-7836(00)00249-6
Rigler FH (1982) The relation between fisheries management
and limnology. Trans Am Fish Soc 111:121–132. doi
:10.1577/1548-8659(1982)111\121:TRBFMA[2.0.CO;2
Roemmich D, McGowan J (1995) Climate warming and the
decline of zooplankton in the California current. Science
267:1324–1326. doi:10.1126/science.267.5202.1324
Rowell TW, Trites RW (1985) Distribution of larval and
juvenile Illex in the Blake Plateau Region (Northwest
Atlantic). Vie Milieu Ser C 35:149–161
Royer TC (1979) On the effect of precipitation and runoff on
coastal circulation in the Gulf of Alaska. J Phys Oceanogr
9:555–563
Rudnick DL, Davis RE (2003) Red noise and regime shifts.
Deep Sea Res Part I Oceanogr Res Pap 50:691–699. doi:
10.1016/S0967-0637(03)00053-0
Sardeshmukh PD, Penland C, Newman M (2001) Drifts
induced by multiplicative red noise with application to
climate. Europhys Lett 63:498–504. doi:10.1209/epl/
i2003-00550-y
Schwager M, Johst K, Jeltsch F (2006) Does red noise increase
or decrease extinction risk? Single extreme events versus
series of unfavorable conditions. Am Nat 167:879–888
Shelton PA, Stansbury DE, Murphy GR et al (–1996) An
assessment of the cod stock in NAFO divisions 2J?3KL.
NAFO SCR Doc. 90/96
Soutar A, Isaacs JD (1974) Abundance of pelagic fish during
the 19th and 20th centuries as recorded in anaerobic
sediments off the California. Fish Bull (Wash D C)
72:257–273
Steele JH (1996) Regime shifts in fisheries management. Fish
Res 2:19–23. doi:10.1016/0165-7836(95)00440-8
Steele JH (1998) Regime shifts in marine ecosystems. Ecol
Appl 8(Suppl):33–36
Steele JH (2004) Regime shifts in the ocean: reconciling
observations and theory. Prog Ocean 60:135–141
Steele JH, Henderson EW (1984) Modeling long-term fluctu-
ations in fish stocks. Science 224:985–987. doi:10.1126/
science.224.4652.985
Stricherz V (1999) Evidence mounts for Arctic oscillation’s
impact on northern climate. http://www.washington.edu/
newsroom/news/1999archive/12-99archive/k121699.html
Thompson JD (1991) The biology of an invasive plant. Bio-
science 41:393–401. doi:10.2307/1311746
191
Torrence C, Compo GP (1998) A practical guide to wavelet
analysis. Bull Am Meteorol Soc 79:61–78. doi :10.1175/
1520-0477(1998)079\0061:APGTWA[2.0.CO;2
Vasseur DA, Yodzis P (2004) The color of environmental
noise. Ecology 85:1146–1152
Wada T, Jacobson LD (1998) Regimes and stock-recruitment
relationships in Japanese sardine (Sardinops melanostic-
tus), 1951–1995. Can J Fish Aquat Sci 55:2455–2463. doi:
10.1139/cjfas-55-11-2455
Ware DM, McFarlane GA (1989) Fisheries production
domains in the northeast Pacific ocean. In: Beamish RJ,
McFarlane GA (eds) Effects of ocean variability on
recruitment and an evaluation of parameters used in stock
assessment models. Can J Fish Aquat Sci (Spec Publ)
108:359–379
Ware DM, Thomson RE (1991) Link between long-term var-
iability in upwelling and fish production in the northeast
Pacific ocean. Can J Fish Aquat Sci 48:2296–2306. doi:
10.1139/f91-270
Watanabe M, Nitta T (1999) Decadal changes in the atmo-
spheric circulation and associated surface climate
variations in the Northern Hemisphere winter. J Clim
12:494–510. doi :10.1175/1520-0442(1999)012\0494:
DCITAC[2.0.CO;2
Werner FE, Perry RI, Lough RG, Naimie CE (1996) Tropho-
dynamic and advective influences on Georges Bank larval
cod and haddock. Deep Sea Res Part II Top Stud Ocea-
nogr 43:1793–1822. doi:10.1016/S0967-0645(96)00042-2
Werner FE, Quinlan JA, Blanton BO et al (1997) The role of
hydrodynamics in explaining variability in fish popula-
tions. J Sea Res 37:195–212. doi:10.1016/S1385-1101
(97)00024-5
Werner FE, Quinlan JA, Lough RG et al (2001) Spatially-
explicit individual based modeling of marine populations:
a review of the advances in the 1990s. Sarsia 86:411–421
Wickett WP (1967) Ekman transport and zooplankton con-
centration in the North Pacific Ocean. J Fish Res Board
Can 24:581–594
Wiebe PH Beardsley RC, Bucklin AC et al (eds) (2001) North
Atlantic GLOBEC studies. Deep Sea Res Part II Top Stud
Oceanogr 48(1–3):1–2
Williamson AH (1992) Historical lobster landings for Atlantic
Canada, 1892–1989. Can Manuscr Rep Fish Aquat Sci
2164: 110pp
Wolfe DA (1985) Fossile fuels: transportation and marine
pollution. Wastes in the ocean 4. In: Duedall IW, Kester
DR, Park PH, Ketchum BH (eds) Energy wastes in the
Ocean. Wiley, New York, pp 45–93
Zhang Y, Wallace JM, Battisti DS (1997) Enso-like inter-
decadal variability: 1900–93. J Clim 10:1004–1020. doi
:10.1175/1520-0442(1997)010\1004:ELIV[2.0.CO;2
Zhang CI, Lee JB, Kim S et al (2000) Climate regime shifts
and their impacts on marine ecosystem and fisheries
resources in Korean waters. Prog Oceanogr 47:171–190.
doi:10.1016/S0079-6611(00)00035-5
123
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