Fisheries Research 55 (2002) 1–9
Viewpoint
Overfishing, tropicalization of fish stocks, uncertainty and
ecosystem management: resharpening Ockham’s razor
Konstantinos I. Stergiou*
Laboratory of Ichthyology, Department of Zoology, School of Biology, Aristotle University of Thessaloniki,
P.O. Box 134, 54006 Thessaloniki, Greece
Received 4 December 2000; accepted 4 December 2000
Abstract
Fishing at the early stage of fisheries development most probably approximated natural predation. Nowadays, fishing
approximates ‘‘extermination’’ with dramatic effects on aquatic ecosystems. Conventional fisheries models and management
practices are inadequate to handle the present situation because, among many other factors, fishing: (a) drives fish stocks to
exhibit smaller body sizes and age/length at maturity (i.e., ‘‘tropicalization’’) and (b) increases catch variability, thus
increasing uncertainty. The realization of (a) and (b) renders conventional practices even more inadequate, thus producing a
never-ending positive feedback loop. Although the application of the precautionary approach to fisheries management together
with the development of indicators and reference point values that trigger management actions seem to be an important step
forward, their adoption within the framework of the same conventional models used to assess fish stocks could introduce
another degree of complexity into existing models. With Ockham’s razor as a primary guiding principle, the advantage of
using ever more complex models is suspect. Ecosystem management seems the only alternative. Within this framework,
alternative simpler ‘‘models and strategies’’ such as large-scale marine protected areas, in which no fishing takes place, are
available and promising, and their adoption as a primary management tool satisfies simultaneously all objectives that have
been set for ecosystem management.
Keywords: Overfishing; Life-history; Tropicalization of fish stocks; Uncertainty; Catch variability; Stock assessment; Precautionary approach;
Ecosystem management
‘‘Fifty years ago, a single cod was large enough
to feed a family of four or five. Today, it is barely
enough for one’’, says Lord Perry of Walton, a
member of the UK House of Lords (Anonymous,
1997b).
There is a worldwide increasing concern for the
future of fisheries resources and their management
mainly because of recent failures in fisheries management and crisis at various levels. Such a concern has
*
Tel.: þ30-31-998268; fax: þ30-31-998279.
E-mail address: kstergio@bio.auth.gr (K.I. Stergiou).
0165-7836/02/$ – see front matter
PII: S 0 1 6 5 - 7 8 3 6 ( 0 1 ) 0 0 2 7 9 - X
been the subject of many recent publications (e.g.,
Mooney, 1998; Pitcher et al., 1998; Flaaten et al.,
1998; Payne, 1999; Briand, 1999; Hollingworth,
2000). In this essay, I tackle issues, albeit not new,
from a different perspective. Thus, I briefly discuss
how fishing activity has changed from natural predation, in early times, to ‘‘extermination’’ today, the
latter being accompanied by important effects on
marine ecosystems. Consequently, I consider in more
depth some fishing-induced effects (i.e., ‘‘tropicalization’’ of fish stocks, increase in objective uncertainty),
which, together with many other facts, contribute,
2
K.I. Stergiou / Fisheries Research 55 (2002) 1–9
through a never-ending loop, to the failure of conventional fisheries models and management practices to
fulfill the very cause of their existence. I further
attempt to show why the implementation of the precautionary approach and the development of ‘‘stockoriented’’ indicators and reference points are not
consistent with the principle of Ockham’s razor and
will not be successful. Finally, I argue that placing
fisheries management into an ecosystem perspective is
the only alternative. Within this framework, the adoption of large-scale, marine protected areas—where
fishing is not allowed—as the primary management
tool satisfies all objectives for ecosystem management, being also consistent with Ockham’s razor.
1. Fishing: from predation to extermination
Fishing was developed gradually when humans
moved from random collection of things found in
nature to the first cultivation and systematic exploitation of food resources using learned practices and
simple tools (Sahrhage and Lundbeck, 1992). Thus,
although about 100,000 years BP Neanderthal man
practiced fishing by hand, by about 50,000 years BP,
Homo sapiens started to use various gears, made of
wood, bone, ivory and horn (Sahrhage and Lundbeck,
1992). By Homer’s time (800–900 B.C.), fishing tools
were already technologically improved (i.e., fine and
pointed curved hooks, made of metal, bronze and iron;
harpoons; nets of mesh sizes of 5–45 mm made of
hemp and flax) (Sahrhage and Lundbeck, 1992).
During the past several centuries, however, the
mechanization of fishing and other developments
(e.g., technological innovations in vessel design,
development of trading organizations and of transport
facilities), allowed fishing activities to expand spatially (Sahrhage and Lundbeck, 1992). Later technological innovations in vessel construction, in fishing
gear material and manufacture, and in electronics
(e.g., radar, echo sounders), the use of highly sizeunselective and efficient gears (e.g., trawls) and the
subsidy-driven overcapitalization of the fishing industry (Beddington, 1995; Anonymous, 1997a; Garcia
and Newton, 1997) all in a synergetic fashion allowed
fishing effort and fishing efficiency to reach unsustainable levels. This has been particularly evident during
the last 10–20 years, with the cost of fishing already
exceeding the value of the world’s catch in the beginning of the 1990s (Cochrane, 2000).
Indeed, very few areas of the world oceans are not
being drastically affected today by fishing activities.
Thus, fisheries landings, together with by-catch,
require about 24–35% of the global marine primary
production in the continental shelf and major upwelling areas of the world ocean, a figure much higher
than previously thought (Pauly and Christensen,
1995). Moreover, 60% of the world fisheries stocks
are fully exploited to overexploited and 6% are
depleted (Anonymous, 1997a), with the case of the
Canadian cod (Gadus morhua) being considered as
one of the biggest disasters of recent years (Spurgeon,
1997a; Longhurst, 1998), and with so far very little
evidence for rapid recovery of stocks from prolonged
declines (Hutchings, 2000). In addition, the mean
trophic level in fisheries landings—i.e., mean trophic
level of all species making up the landings weighted
by their catch—in the last 45 years has decreased
steadily both at the global scale and at the regional,
ocean-specific scale (i.e., the fishing-down the food
web concept: Pauly et al., 1998a,b, 2000a). Finally,
fishing-induced species’ extinctions, or near extinctions, seem to be more frequent than previously
thought (see review by Roberts and Hawkins,
1999), even though fisheries models do predict them
(Pitcher, 1998).
Thus, humans, at the early stage of fisheries development, used fishing gears (e.g., nets, hooks, harpoons) that we now know are highly size-selective.
Since their effect on aquatic ecosystems was spatially
very localized, one may conclude that fishing during
this early period most probably approximated natural
predation. In contrast, there is no doubt that nowadays
fishing approximates ‘‘extermination’’, with dramatic
effects on aquatic ecosystems and important implications for fisheries management both of which are
discussed below.
2. Tropicalization of fish stocks and objective
uncertainty: the epitaph of conventional fisheries
models and management schemes
The ecosystem effects of fishing, some of which
have been described above, can be generally classified
into two categories (Fig. 1). The first includes effects
K.I. Stergiou / Fisheries Research 55 (2002) 1–9
3
Fig. 1. A schematic, simplified representation of the major effects of fishing on marine ecosystems (modified from Stergiou, 1999). Key or
example references per major effect portrayed on this graph are given below. Effects at the community level: (a) destruction of structure and
heterogeneity of benthic habitats (see review by Jennings and Kaiser, 1998); (b) changes in relative species composition (e.g., Pauly, 1994a;
Caddy and Rodhouse, 1998; Fogarty and Murawski, 1998; Stevens et al., 2000); (c) decrease in species diversity (e.g., Rijnsdorp et al., 1996;
Stergiou et al., 1997a; Bianchi et al., 2000; but see Greenstreet and Hall, 1996); (d) species’ extinction (see review by Roberts and Hawkins,
1999); (e) changes in predation and competition rates, trophic cascades (e.g., Roberts and Polunin, 1991; Ramsay and Kaiser, 1998; Sala et al.,
1998); (f) changes in trophic structures and energy flow (Pauly and Christensen, 1995, 1998a,b, 2000a); (g) decrease in stock abundances (e.g.,
Haedrich and Barnes, 1997; Anonymous, 1997a; Hutchings, 2000; Clark et al., 2000); (h) increase in variability with time (Stergiou, 1998).
Effects of fishing at the life-history level of the individual species: (a) decrease in body size/weight (e.g., McAllister et al., 1992; Haedrich and
Barnes, 1997; Ratz et al., 1999; Bianchi et al., 2000; Zwanenburg, 2000); (b) size and age at maturity: (Beacham, 1983; Jennings and Kaiser,
1998; Morgan and Colbourne, 1999; Law, 2000); (c) truncating age structures (e.g., Longhurst, 1998); (d) change in sex ratios (e.g., Buxton,
1993); and (e) decrease in population reproductive potential (e.g., Jennings and Kaiser, 1998; Jennings et al., 1998).
at the community level and the second those at the lifehistory level of the individual species. There is no
doubt that the effects at the two levels, which can be
either direct or indirect (e.g., Gislason et al., 2000),
strongly interact with each other in a complex and
often unpredictable manner.
From the various effects of fishing, I hereafter pay
particular attention to those concerning the length
structure and the length/age at maturity of the
exploited stocks. The examples presented in Table 1
suffice to make the point that such effects, which may
have a genetic basis (e.g., Policansky and Magnuson,
1998; Law, 2000), can be very dramatic indeed. They
also clearly show that heavy fishing in non-tropical
waters ‘‘tropicalizes’’ fish stocks, in the sense that it
drives them to exhibit the characteristics of their
tropical counterparts (i.e., smaller body sizes, earlier
maturation: Pauly, 1998a). Such a ‘‘tropicalization’’
renders the application of the presently used conventional stock assessment models inadequate (e.g., Die
4
K.I. Stergiou / Fisheries Research 55 (2002) 1–9
Table 1
Examples of the effects of fishing on body size and length/age at maturity in fishes
Species
Effect of fishing
Reference
Various species
Based on data from FishBase 1998, the mean maximum length of the fishes
making up the catches of the NE and NW Atlantic has decreased from about
90 cm in 1950 to 60 cm in 1996
The mean weight of commercially targeted demersal fishes declined by 51
and 41% on the eastern and western Scotian shelf, respectively, between
1970–1974 and 1994–1998; in both areas the decline was faster during
mid-1980s to mid-1990s
The mean weight of adults has been reduced by up to 34% for several
British Columbia stocks between 1950 and 1990
The length-at-age 4 and 5 years declined from about 61 and 67 cm in
1959 to 43 and 53 cm in the mid-1980s, respectively
The mean weight of the northern Grand Banks (NAFO Div. 2J3KL) cod
declined from >2 kg to about 0.5 kg after the collapse of the early 1990s,
with the biomass of the 10–20-year-old individuals being reduced from
48% in 1962 to 8% in 1990 and to zero after 1990
The median length and age at maturity of the majority of the stocks on the
Scotian shelf, which were heavily exploited in the 1960s and 1970s,
declined by about 50% between 1959 and 1979 (from about 6 to
3 years and from 55 cm to about less than 45 cm
The median age at maturity of the Northeast cod declined drastically from
11 years in the late 1940s to less than 8 years in the late 1980s
The age and length at 50% maturity of the three major stocks,
the Labrador-NE Newfoundland, Grand Bank and St Pierre Bank stocks,
have both declined drastically between early 1960s and late 1980s
(by 2–5 years and 3–10 cm depending on stock and sex)
Froese and Pauly (1998)
Various species
Oncorhynchus gorbuscha
Gadus morhua
Gadus morhua
Gadus morhua
Gadus morhua
Hippoglossoides platessoides
and Caddy, 1997) and calls for the use of their
‘‘tropicalized’’ versions (see Pauly, 1998a).
Body length is the most important demographic
characteristic of a species that largely determines
many processes: ecological (e.g., predation, competition, trophic level, mortality, longevity: Pauly,
1998a,b; variability: Stergiou, 1998); fisheries (e.g.,
gear size selectivity: Pauly, 1998a); and managerial
(e.g., risk undertaken by fishery managers: Stergiou,
1998). Based on FAO catch data from 103 fish stocks
throughout the world oceans, Stergiou (1998; unpublished data) showed that catch variability decreases
with an increase in body size. Because however, there
is a strong, significant relationship between body
length and trophic level (Froese and Pauly, 1998),
one may assume that fishing-down marine food webs,
by decreasing body sizes (Pauly et al., 1998b) and
truncating age structures, should increase catch variability. Indeed, Stergiou (1998) has also shown that the
variability of fish catches during the last years
increases with the length of time over which it is
Zwanenburg (2000)
McAllister et al. (1992)
Ratz et al. (1999)
Longhurst (1998)
Beacham (1983)
Jorgensen (1990)
Morgan and Colbourne
(1999)
calculated (Fig. 2; sensu Pimm and Redfearn, 1988).
This indicates that catches exhibit long-term trends
and hence that there is not any equilibrium yield, the
latter being the basis of the conventional single- and
multi-species models presently used for fisheries management (e.g., Caddy and Sharp, 1986; Hilborn and
Walters, 1992; Hollowed et al., 2000).
Temporal variability in catches, and other parameters such as recruitment, introduce what is usually
referred to as ‘‘objective uncertainty’’ (Ferson and
Ginzburg, 1996). Objective uncertainty refers to
uncertainties resulting from the underlying variability
in stochastic processes, such as growth, mortality and
recruitment, as opposed to ‘subjective uncertainty’
(Ferson and Ginzburg, 1996), which is mainly attributed to lack of knowledge (e.g., the case for many
Eastern Mediterranean stocks: Stergiou et al., 1997b).
Although it is widely recognized that objective uncertainty is one of the key-factors in predicting and
managing fisheries resources (e.g., Walters and
Maguire, 1996; Flaaten et al., 1998; Stokes et al.,
K.I. Stergiou / Fisheries Research 55 (2002) 1–9
Fig. 2. Variability of catches (standard deviation of logged catches,
SDL) versus the number of years over which SDL was calculated
for Mallotus villosus in Norwegian waters for the period 1918–
1991 (data from Stergiou (1984)). SDL is plotted against 2, 4, 8,
16, 32, 64 and 72 years. The slope of the regression line is
significantly ðP < 0:05Þ different from zero (from Stergiou, 1998).
1999; Cochrane, 2000), and managers are aware of
such uncertainty, it is still largely neglected in stock
assessment or management procedures (Lauck et al.,
1998). In fact, it is to this aspect of fisheries ecology
that recent paradigms of fish stocks collapses are
generally attributed to (e.g., Anonymous, 1995; Spurgeon, 1997b). Both, the fishing-induced tropicalization of fish stocks and the increase in uncertainty are
involved in a never-ending positive loop (i.e., they will
further contribute to the inadequacy of models and
management strategies, which, in turn, will intensify
tropicalization and uncertainty).
Another point to be considered here is the effect of
climate on fisheries, which further complicates things
for fisheries scientists and managers. This is because
most of the effects of fishing on ecosystems and
individual species (Fig. 1) can also be brought about
by, or related to, long-term climatic changes, as indicated by several earlier (e.g., Jensen, 1939; Dunbar,
1954; Cushing and Dickson, 1976) and recent examples of the effect of climate on marine communities
and stocks (see Stergiou, 1999). In fact some of the
effects shown in Table 1 have been partially related to
climate (e.g., Ratz et al., 1999). The same may also be
true of various anthropogenic impacts such as pollution and habitat modification (e.g., Caddy, 1993;
Gislason et al., 2000). The case of ‘‘regime shifts’’
is also relevant here. Regime shifts are synchronized
5
changes in several commercial stocks at long, interdecadal, scales and these changes, being produced or
enhanced by climatic changes either natural or anthropogenic ones, can be linked to changes in other
components of marine ecosystems such as the abundance of planktonic invertebrates and temperature
(Bakun, 1998; Steele, 1998). Steele (1998) maintains
that because of the ‘‘regime shifts’’, the ‘‘sustainability’’ concept is not directly applicable to marine
ecosystems and especially to fisheries. It must also
be kept in mind that the effect of the climate and of the
environment cannot be distinguished from that of
fishing inasmuch as fisheries managers will always
respond to catch declines by assuming that fishing is
the main factor and, hence, both effects will be
reflected in catch records (Hilborn and Walters, 1992).
Thus, conventional fisheries models are inadequate
for dealing with the present situation because of the:
(a) fishing-induced tropicalization of fish stocks; (b)
fishing-induced increase in objective uncertainty; (c)
increase in catch variability with the time over which it
is estimated (i.e., non-equilibrium yield); and (d) the
confounding effects of climate and environment on
fisheries stocks, which are not generally accounted for
in the models.
The above mentioned issues together with the facts
that: (a) actual fishing mortality rates usually exceed
target rates (e.g., because of unreported catches, discarding, ghost fishing: Lauck et al., 1998), (b) enforcement of traditional measures, such as limits by
numbers, minimum landings sizes, is very difficult
(Anonymous, 1997b; Stergiou et al., 1997a), (c) managerial decisions are, and will be, highly influenced by
politics (Masood, 1997; Spurgeon, 1997b,c), and (d)
objectives are poorly articulated in fisheries managements schemes (Stokes et al., 1999; Cochrane, 2000),
have all, in a synergetic fashion, contributed to the
recent failure of conventional fisheries management to
fulfill the very cause of its existence (for more on the
failure of management practices see: Smith, 1998;
Cochrane, 2000).
Thus, conventional fisheries models and management strategies should be reconsidered in order to
account for the above issues. Recently, the ‘‘precautionary approach’’ has been applied to fisheries management, while the idea of ‘‘ecosystem management’’
has also been put on the table. These issues are
discussed below.
6
K.I. Stergiou / Fisheries Research 55 (2002) 1–9
3. The precautionary approach, ecosystem
management and Ockham’s razor
A version of the precautionary approach derived
from Principle 15 of the Rio Declaration reads:
‘‘Where there are threats of serious or irreparable
damage, lack of full scientific certainty shall not be
used as a reason for postponing cost-effective measures
to prevent environmental degradation’’. The application of this principle to fisheries management and the
development of sustainability indicators (e.g., stock
biomass, fishing mortality) in relation to reference
point values that trigger management actions for the
target species (e.g., Garcia, 1994; Darcy and Matlock,
1999; Oliver, 2000), seems to be an important step
forward. However, the adoption of ‘‘stock-oriented’’
indicators and reference points will be unsuccessful
because they are implemented within the framework of
the same conventional models used to assess fish
stocks. In addition, the use of single-species reference
points is questionable when species interactions are
important (Gislason, 1999), as is usually the case for
most fisheries. The adoption of ‘‘stock-oriented’’ indicators and reference points could also introduce
another degree of complexity into the existing models
(see for instance Gislason, 1999), with doubtful outcomes. It is noteworthy that the incorporation of longterm trends and cycles in commercial catches (e.g.,
Taylor and Prochaska, 1984), of uncertainties (e.g.,
Punt and Hilborn, 1997), or environmental factors
(e.g., Basson, 1999) into various single-species stock
assessment models, also adds complexity with doubtful outcomes. Indeed, attempts to address uncertainties
generally lead to increased complexity in management
systems without any sign of having the desired results,
even for the commercial single-species fisheries for
which they were developed (Cochrane, 1999). Thus,
the adoption of ‘‘stock-oriented’’ reference points will
delay the identification of the detrimental effects
of fishing on any given stock, a fact that by itself
may further contribute to the failure of management
practices (Lauck et al., 1998).
Placing fisheries management into an ecosystem
perspective seems to be the only alternative (e.g.,
Jennings and Polunin, 1996; Mooney, 1998; Gislason
et al., 2000; Cochrane, 2000; Pitcher, 2000), and the
adoption of a variety of ‘‘ecosystem’’ indicators and
reference points that trigger management actions
(e.g., Table 1 in Gislason et al., 2000) becomes a
necessity. These can be defined directly, through the
use of modeling tools such as the ECOPATH/ECOSIM/ECOSPACE (Christensen and Pauly, 1993a;
Walters et al., 1997; Pauly et al., 2000b), which are
invaluable for studying the effects of fisheries, and
have been successfully applied in various areas (e.g.,
Christensen and Pauly, 1993b; Shannon et al., 2000).
Ecosystem objectives may also be satisfied indirectly, by establishing, what is considered to be the
extreme case of the precautionary approach (Lauck
et al., 1998), marine protected areas (MPAs; also
known as no-take zones). MPAs, in which fishing is
totally prohibited, provide a refuge in space rather than
a refuge in numbers, the latter being the aim of most
traditional fisheries management measures. Among
many other benefits, MPAs protect the biomass of
species, maintain biodiversity including genetic diversity, decrease the trend for heavy evolutionary fishing
selection for earlier maturity and reproduction and
smaller adult fish size, and hedge against inevitable
uncertainties, errors in estimations, and biases (e.g.,
Roberts and Polunin, 1991; Agardy, 1994; Lauck et al.,
1998; Guenette et al., 1998; Hall, 1998; Sumaila et al.,
2000; Polunin, 2001). Thus MPAs satisfy simultaneously the objectives for ecosystem management
proposed by Gislason et al. (2000) (maintenance of:
ecosystem diversity, species diversity, within-species
genetic variability, directly impacted species, ecologically dependent species and trophic level balance) or
others (see Fowler, 1999). However, in the large-scale
ecological context, the ability of MPAs to meet such
objectives depends critically, apart from the degree of
enforcement, on their size and the biology and mobility of the target species concerned (see thorough
review by: Polunin, 2001). Such dependence might
be true for the vast majority of the existing MPAs,
which are of very small sizes (i.e., their median size
being 1600 ha). In fact, such sizes do not always have
fisheries or ecosystem benefits (Polunin, 2001). Yet,
there is overwhelming evidence that MPA sizes
encompassing large parts, e.g., 40% or more, of the
fishable management areas (e.g., Sladek Nowlis and
Roberts, 1999; Polunin, 2001) could have substantial
benefits in rebuilding highly overfished stocks and
thus the ecosystems in which they are embedded.
I now turn to consider Ockham’s razor (e.g., Pauly,
1994b). With this principle as a primary guide, I
K.I. Stergiou / Fisheries Research 55 (2002) 1–9
wonder what is the advantage of using ever more
complex models, and management strategies based
upon them (i.e., single- or multi-species fisheries models further complicated by reference points), which are
likely to fail when alternative simpler ‘‘models and
strategies’’ such as large-scale MPAs (see Agardy,
2000) are available and promising?
The establishment of effectively large-sized MPAs
must proceed quickly (Ludwig et al., 1993; Fogarty,
1999) and everybody that was previously dependent
on these large areas must be provided with alternative
sources of income. Discussions on issues such as the
identification of optimum MPA size and location,
socio-economic and political repercussions, and many
others, albeit very important at some later stage, serve
at present only as excuses for not adopting MPAs as a
primary management tool. Is there anybody thinking
that the establishment of an MPA will be ecologically
harmful within the boundaries of the ecosystem where
this was established? (It may, over the short term, be
economically harmful, which is all that politicians and
fishermen care about). Should large scale MPAs not be
adopted as a primary management tool, we will
‘‘watch’’ more stocks collapse and maybe then use
what will be left of natural fisheries to support an
aquaculture production of fish and shellfish.
Acknowledgements
I would like to extend my gratitude to Drs. H.
Browman, K. Erzini and A.D. McIntyre for their
constructive criticisms and comments.
References
Agardy, T., 1994. Advances in marine conservation: the role of
marine protected areas. Trends Ecol. Evol. 9, 267–270.
Agardy, T., 2000. Effects of fisheries on marine ecosystems:
a conservationist’s perspective. ICES J. Mar. Sci. 57, 761–
765.
Anonymous, 1995. The cod that disappeared. New Scientist,
September, 1995, 24–29.
Anonymous, 1997a. Review of the state of world fishery resources:
marine fisheries. FAO Fish. Circ. 920, 173.
Anonymous, 1997b. Fishing by numbers reveals its limits. Nature
386, 110.
Bakun, A., 1998. Ocean triads and radical interdecadal variation:
bane and boon to scientific fisheries management. In: Pitcher,
7
T., Hart, P.J., Pauly, D. (Eds.), Reinventing Fisheries Management. Chapman & Hall, London, pp. 332–358.
Basson, M., 1999. The importance of environmental factors in the
design of management procedures. ICES J. Mar. Sci. 56, 933–
942.
Beacham, T.D., 1983. Variability in median size and age at sexual
maturity of Atlantic cod, Gadus morhua, on the Scotian shelf in
the Northwest Atlantic Ocean. Fish. Bull. US 81, 303–321.
Beddington, J., 1995. The primary requirements. Nature 374, 213–
214.
Bianchi, G., Gislason, H., Graham, K., Hill, L., Jin, X., Koranteng,
K., Manickchand-Heilman, S., Paya, I., Sainsbury, K., Sanchez,
F., Zwanenburg, K., 2000. Impact of fishing on size composition and diversity of demersal fish communities. ICES J. Mar.
Sci. 57, 558–571.
Briand, F. (Ed.), 1999. Precautionary Approach to Local Fisheries
in the Mediterranean Sea. CIESM Workshop Series 7, 89 pp.
Buxton, C.D., 1993. Life-history changes in exploited reef fishes on
the east coast of South Africa. Environ. Biol. Fish. 36, 47–63.
Caddy, J.F., 1993. Toward a comparative evaluation of human
impacts on fishery ecosystems of enclosed and semi-enclosed
seas. Rev. Fish. Sci. 1, 57–95.
Caddy, J.F., Rodhouse, P.G., 1998. Cephalopod and groundfish
landings: evidence for ecological change in global fisheries?
Rev. Fish Biol. Fish. 8, 431–444.
Caddy, J.F., Sharp, G.D., 1986. An ecological framework for
marine fishery investigations. FAO Fish. Tech. Pap. 283, 152.
Christensen, V., Pauly, D., 1993a. ECOPATH II, a software for
balancing steady-state ecosystem models and calculating
network characteristics. Ecol. Model. 61, 169–185.
Christensen, V., Pauly, D. (Eds.), 1993b. Trophic Models of
Aquatic Ecosystems. Proceedings of the ICLARM Conference,
Vol. 26, 390 pp.
Clark, M.R., Owen, F.A., Francis, R.I.C., Tracey, D.M., 2000. The
effects of commercial exploitation on orange roughy (Hoplostethus atlanticus) from the continental slope of the Chatham
Rise, New Zealand, from 1979 to 1997. Fish. Res. 45, 217–238.
Cochrane, K.L., 1999. Complexity in fisheries and limitations in
the increasing complexity of fisheries management. ICES J.
Mar. Sci. 56, 917–926.
Cochrane, K.L., 2000. Reconciling sustainability, economic
efficiency and equity in fisheries: the one that got away? Fish
Fish. 1, 3–21.
Cushing, D.H., Dickson, R.R., 1976. The biological responses in
the sea to climatic changes. Adv. Mar. Biol. 14, 1–122.
Darcy, G.H., Matlock, G.C., 1999. Application of the precautionary
approach in the national standard guidelines for conservation
and management of fisheries in the United States. ICES J. Mar.
Sci. 56, 853–859.
Die, D.J., Caddy, J.F., 1997. Sustainable indicators from biomass:
are there appropriate reference points for use in tropical
fisheries? Fish. Res. 32, 69–79.
Dunbar, M.J., 1954. A note on climatic change in the sea. Arctic 7,
27–30.
Ferson, S., Ginzburg, L.R., 1996. Different methods are needed to
propagate ignorance and variability. Reliab. Engr. Syst. Saf. 54,
133–144.
8
K.I. Stergiou / Fisheries Research 55 (2002) 1–9
Flaaten, O., Salvanes, A.G.V., Scweder, T., Ultang, O. (Eds.), 1998.
Objectives and Uncertainties in Fisheries Management with
Emphasis on Three North Atlantic Ecosystems: A Selection of
Papers Presented at an International Symposium, Bergen,
Norway, June 3–5, 1997. Fish. Res. 37, 1–310.
Fogarty, M.J., 1999. Essential habitat, marine reserves and fishery
management. Trends Ecol. Evol. 14, 133–134.
Fogarty, M.J., Murawski, S.A., 1998. Large-scale disturbance and
the structure of marine systems: fishery impacts on Georges
Bank. Ecol. Appl. 8 (Suppl.), 6–22.
Fowler, C.W., 1999. Management of multi-species fisheries: from
overfishing to sustainability. ICES J. Mar. Sci. 56, 927–932.
Froese, R., Pauly, D. (Eds.), 1998. Fishbase 98: Concepts, Design
and Data Sources. ICLARM, Manila.
Garcia, S.M., 1994. The precautionary principle: its implications in
capture fisheries management. Ocean Coast. Mgmt. 22, 99–125.
Garcia, S.M., Newton, C., 1997. Current situation trend and
prospects in world capture fisheries. In: Pikitch, E., Huppert,
D.D., Sissenwine, M. (Eds.), Global Trends: Fisheries Management. American Fisheries Society Symposium, Vol. 20,
Bethesda, USA, pp. 3–27.
Gislason, H., 1999. Single and multispecies reference points for
Baltic fish stocks. ICES J. Mar. Sci. 56, 571–583.
Gislason, H., Sinclair, M., Sainsbury, K., O’Boyle, R., 2000.
Symposium overview: incorporating ecosystem objectives
within fisheries management. ICES J. Mar. Sci. 57, 468–475.
Greenstreet, S.P.R., Hall, S.J., 1996. Fishing and the ground-fish
assemblage structure in the North-western North Sea: an analysis
of long-term and spatial trends. J. Anim. Ecol. 65, 577–598.
Guenette, S., Lauck, T., Clark, C., 1998. Marine reserves: from
Beverton and Holt to the present. Rev. Fish Biol. Fish. 8, 251–
272.
Haedrich, R.L., Barnes, S.M., 1997. Changes over time of the size
structure in an exploited shelf fish community. Fish. Res. 31,
229–239.
Hall, S.J., 1998. Closed areas for fisheries management—the case
consolidates. Trends Ecol. Evol. 13, 297–298.
Hilborn, R., Walters, C.J., 1992. Quantitative Fisheries Stock
Assessment: Choice, Dynamics and Uncertainty. Chapman &
Hall, New York, 570 pp.
Hollingworth, C.E. (Ed.), 2000. Ecosystem Effects of Fishing.
Proceedings of the ICES/SCOR Symposium. ICES J. Mar. Sci.
57, 465–791.
Hollowed, A.B., Bax, N., Beamish, R., Collie, J., Fogarty, M.,
Livingston, P., Pope, J., Rice, J.C., 2000. Are multispecies
models an improvement on single-species models for measuring fishing impacts on marine ecosystems? ICES J. Mar. Sci.
57, 707–719.
Hutchings, J.A., 2000. Collapse and recovery of marine fishes.
Nature 406, 882–885.
Jennings, S., Kaiser, M.J., 1998. The effects of fishing on marine
ecosystems. Adv. Mar. Biol. 34, 201–352.
Jennings, S., Polunin, V.C., 1996. Impacts of fishing on tropical
reef ecosystems. Ambio 25, 44–49.
Jennings, S., Reynolds, J.D., Mills, S.C., 1998. Life history
correlates of responses to fisheries exploitation. Proc. R. Soc.
London B 265, 333–339.
Jensen, A.S., 1939. Concerning a change of climate during recent
decades in the Arctic and sub-Arctic regions, from Greenland
in the west to Eurasia in the east, and countemporary biological
and geophysical changes. Biol. Mediterr. 14, 1–83.
Jorgensen, T., 1990. Long-term changes in age at sexual maturity
of Northeast Arctic cod (Gadus morhua L.). J. Cons. Int.
Explor. Mer. 46, 235–248.
Lauck, T., Clark, C., Mangel, M., Munro, G.R., 1998. Implementing the precautionary principle in fisheries management
through marine reserves. Ecol. Appl. 8, 72–78.
Law, R., 2000. Fishing, selection, and phenotypic evolution. ICES
J. Mar. Sci. 57, 659–668.
Longhurst, A., 1998. Cod: perhaps if we all stood back a bit? Fish.
Res. 38, 101–108.
Ludwig, D., Hilborn, R., Walters, C., 1993. Uncertainty, resource
exploitation, and conservation: lessons from history. Science
260, 36.
Masood, E., 1997. Fisheries science: all at sea when it comes to
politics? Nature 386, 105–106.
McAllister, M.K., Peterman, R.M., Gillis, D.M., 1992. Statistical
evaluation of a large scale fishing experiment designed to test
for a genetic effect of size-selective fishing on British Columbia
pink salmon (Oncorhynchus gorbuscha). Can. J. Fish. Aquat.
Sci. 49, 1294–1304.
Mooney, H.A. (Ed.), 1998. Ecosystem Management for Sustainable
Marine Fisheries. Ecol. Appl. 8 ( Suppl. 1), 1–174.
Morgan, M.J., Colbourne, E.B., 1999. Variation in maturity-at-age
and size in three populations of American plaice. ICES J. Mar.
Sci. 56, 673–688.
Oliver, P., 2000. The knowledge on the state of Mediterranean
resources in relation to their sustainable management within the
precautionary approach to fisheries: recent initiatives and
proposals to fill the gaps. CIESM Workshop Series 12, 53–56.
Pauly, D., 1994a. On the Sex of Fish and the Gender of Scientists:
A Collection of Essays in Fisheries Science. Fish and Fisheries
Series 14. Chapman & Hall, London.
Pauly, D., 1994b. Resharpening Ockham’s razor. Naga 17 (2), 7–8.
Pauly, D., 1998a. Beyond our original horizons: the tropicalization
of Beverton and Holt. Rev. Fish Biol. Fish. 8, 307–334.
Pauly, D., 1998b. Tropical fishes: patterns and propensities. J. Fish.
Biol. 53 (Suppl.), 1–17.
Pauly, D., Christensen, V., 1995. Primary production required to
sustain global fisheries. Nature 374, 255–257.
Pauly, D., Christensen, V., Dalsgaard, J., Froese, R., Torres Jr., F.,
1998a. Fishing down marine food webs. Science 279, 860–
863.
Pauly, D., Froese, R., Christensen, V., 1998b. Response to Caddy
et al. How pervasive is fishing down marine food webs?
Science 282, 1384–1386.
Pauly, D., Christensen, V., Froese, R., Palomares, M.L., 2000a.
Fishing down aquatic food webs. Am. Sci. 88, 46–51.
Pauly, D., Christensen, V., Walters, C., 2000b. Ecopath, ecosim,
and ecospace as tools for evaluating ecosystem impacts on
marine ecosystems. ICES J. Mar. Sci. 57, 697–706.
Payne, A.I.L. (Ed.), 1999. Confronting Uncertainty in the
Evaluation and Implementation of Fisheries-management
Systems. ICES J. Mar. Sci. 56.
K.I. Stergiou / Fisheries Research 55 (2002) 1–9
Pimm, S.L., Redfearn, A., 1988. The variability of population
densities. Nature 334, 613–614.
Pitcher, T.J., 1998. A cover story: fisheries may drive stocks to
extinction. Rev. Fish Biol. Fish. 8, 367–370.
Pitcher, T.J., 2000. Ecosystem goals can reinvigorate fisheries
management, help dispute resolution and encourage public
support. Fish Fish. 1, 99–103.
Pitcher, T.J., Hart, P.J.B., Pauly, D. (Eds.), 1998. Reinventing
Fisheries Management. Chapman & Hall, London.
Policansky, D., Magnuson, J.J., 1998. Genetics, metapopulations,
and ecosystem management of fisheries. Ecol. Appl. 8 (Suppl.),
119–123.
Polunin, N.V.C., 2001. Marine protected areas, fish and fisheries.
In: Hart, P.J.B., Reynolds, J.C. (Eds.), Handbook of Fish and
Fisheries, Vol. II. Blackwell, Oxford.
Punt, A.E., Hilborn, R., 1997. Fisheries stock assessment and
decision analysis: the Bayesian approach. Rev. Fish Biol. Fish.
7, 35–63.
Ramsay, K., Kaiser, M.J., 1998. Demersal fishing disturbance
increases predation risk for whelks (Buccinum undatum L.). J.
Sea Res. 39, 299–304.
Ratz, H.-J., Stein, M., Lloret, J., 1999. Variation in growth and
recruitment of Atlantic cod (Gadus morhua) off Greenland
during the second half of the 20th century. J. Nortw. Atl. Fish.
Sci. 25, 161–170.
Rijnsdorp, A.D., van Leeuwen, P.I., Daan, N., Heesen, H.J.L.,
1996. Changes in abundance of demersal fish species in the
North Sea between 1906–1909 and 1990–1995. ICES J. Mar.
Sci. 53, 1054–1062.
Roberts, C.M., Hawkins, R., 1999. Species extinctions in marine
ecosystems. Trends Ecol. Evol. 14, 241–246.
Roberts, C.M., Polunin, N.V.C., 1991. Are marine reserves effective
in management of reef fisheries? Rev. Fish Biol. Fish. 1, 65–91.
Sahrhage, D., Lundbeck, J., 1992. A History of Fishing. Springer,
Berlin.
Sala, E., Boudouresque, C.F., Harmelin-Vivien, M., 1998. Fishing,
trophic cascades, and the structure of algal assemblages:
evaluation of an old but untested paradigm. Oikos 83, 425–439.
Shannon, L.J., Cury, P.M., Jarre, A., 2000. Modelling effects of
fishing in the southern Benguela ecosystem. ICES J. Mar. Sci.
57, 720–722.
Sladek Nowlis, J.S., Roberts, C.M., 1999. Fisheries benefits and
optimal design of marine reserves. Fish. Bull. US 97, 604–616.
Smith, T.D., 1998. Simultaneous and complementary advances:
mid-century expectations of the interaction of fisheries science
and management. Rev. Fish Biol. Fish. 8, 335–348.
Spurgeon, D., 1997a. Canada’s cod leaves science in hot water.
Nature 386, 107.
9
Spurgeon, D., 1997b. Scientists dispute wisdom of Canada
reopening fishery. Nature 386, 748.
Spurgeon, D., 1997c. Political interference skewed scientific advice
on fish stocks. Nature 388, 106.
Steele, J.H., 1998. Regime shifts in marine ecosystems. Ecol. Appl.
8, 33–36.
Stergiou, K.I., 1984. Capelin and climatic change in the Barents
Sea. M.Sc. Thesis. McGill University, Montreal, Canada, 225
pp.
Stergiou, K.I., 1998. Variability of fish catches in different
ecosystems. In: Durand, M.E., Cury, P., Mendelssohn, R.,
Roy, C., Bakun, A., Pauly, D. (Eds.), Global Versus Local
Changes in Upwelling Systems. ORSTOM Editions, Paris,
pp. 359–370.
Stergiou, K.I., 1999. Precaution in fisheries within the context of
ecological and environmental changes. CIESM Workshop Ser.
7, 33–36.
Stergiou, K.I., Politou, Ch.-Y., Christou, E.D., Petrakis, G., 1997a.
Selectivity experiments in the NE Mediterranean: the effect of
trawl codend mesh size on species diversity and discards. ICES
J. Mar. Sci. 54, 774–786.
Stergiou, K.I., Christou, E., Georgopoulos, D., Zenetos, A.,
Souvermezoglou, C., 1997b. The Hellenic Seas: physics,
chemistry, biology and fisheries. Oceanogr. Mar. Biol. Ann.
Rev. 35, 415–538.
Stevens, J.D., Bonfil, R., Dulvy, N.K., Walker, P.A., 2000. The
effects of fishing on sharks, rays, and chimaeras (chondrichthyans), and the implications for marine ecosystems. ICES J.
Mar. Sci. 57, 476–494.
Stokes, T.K., Butterworth, D.S., Stephenson, R.L., Payne, A.I.L.,
1999. Confronting uncertainty in the evaluation and implementation of fisheries-management systems. Introduction.
ICES J. Mar. Sci. 56, 795–796.
Sumaila, U.R., Guenette, S., Alder, J., Chuenpagdee, R., 2000.
Addressing ecosystem effects of fishing using marine protected
areas. ICES J. Mar. Sci. 57, 752–760.
Taylor, G.T., Prochaska, F.J., 1984. Incorporating unobserved
cyclical stock movements in fishery catch equations: an
application to the Florida blue crab fishery. N. Am. J. Fish.
Mgmt. 4, 67–74.
Walters, C., Maguire, J.J., 1996. Lessons for stock assessment from
the northern cod collapse. Rev. Fish Biol. Fish. 6, 125–137.
Walters, C., Christensen, V., Pauly, D., 1997. Structuring dynamic
models of exploited ecosystems from trophic mass-balance
assessments. Rev. Fish Biol. Fish. 7, 139–172.
Zwanenburg, K.C.T., 2000. The effects of fishing on demersal fish
communities of the Scotian shelf. ICES J. Mar. Sci. 57, 503–
509.