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Aquatic Ecosystems
Trends and Global Prospects
Edited by Nicholas V. C. Polunin
Book DOI: http://dx.doi.org/10.1017/CBO9780511751790
Online ISBN: 9780511751790
Hardback ISBN: 9780521833271
Chapter
20 - The marine pelagic ecosystem: perspectives on humanity's role in the future pp. 311-318
Chapter DOI: http://dx.doi.org/10.1017/CBO9780511751790.028
Cambridge University Press
20 The marine pelagic ecosystem: perspectives on
humanitys role in the future
p e t e r g. v e r i t y, j o h n h. s t e e l e , t. f r e d e t h i n g s ta d a n d
f e r e i d o u n r a s s o u l z a d e g a n
INTRODUCTION
Hensen (1887) proposed that food supply controlled vari-
ations in adult sh stocks, and therefore quantitative studies
of plant and animal production in the sea might permit
predictions of annual sh yields. If sh could be harvested
by man, Hensen (1887) argued that relationships similar to
agriculture existed between primary production and sh
yield. The original conceptual basis for quantitative marine
ecology has a terrestrial origin (Verity et al. 2002). In
retrospect, however, it is now appreciated that natural
vegetation on land, such as that of trees, has long time-scales
compared to marine primary producers (Steele 1991).
Unlike terrestrial systems, marine pelagic communities
appear to be much more adaptable to natural drivers and
anthropogenic pressures, at least at timescales of years to
decades.
The differences between marine and terrestrial sys-
tems are more important than the similarities. These
Aquatic Ecosystems, ed. N. V. C. Polunin. Published by Cambridge University Press. Foundation for Environmental Conservation 2008.
311
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differences include the way that marine systems respond
to change, differences in the ways that humans attempt
management and differences in how the terrestrial con-
cept of sustainability relates to similar practices in the
marine pelagic ecosystem. For example, the long time-
scales of the natural vegetation on land are reected in the
regeneration of forests, which can take decades to cen-
turies. One potential concern with global warming is that
plant communities could not adapt to the relatively rapid
latitudinal shift in temperature and so could not maintain
optimal conditions for growth. This is not really a
problem in the sea, where the base of the food web, the
phytoplankton, has lifetimes of days not decades. Fish,
residing at the top of the marine web, mainly live for a
few years, although some in the deep sea (Chapter 22) and
off coral reefs (Chapter 16) live for decades. These factors
have important implications for the effective application
of scientic understanding to management of all marine
resources and furthermore to human policies that affect
the pelagic ecosystem in other ways, such as pollution,
bioinvasions, diseases and eutrophication.
Humanity evolved in the terrestrial world, which is
therefore intuitively better understood compared to aqua-
tic environments. But human activities, planned and
unplanned, are now causing dramatic changes in the latter,
even systems as physically huge as the marine pelagic
realm. This chapter will provide an overview of the status
of anthropogenic impacts on the marine pelagic ecosystem
by comparison with natural phenomena and, using marine
sheries as an example, suggest strategies for sustainability
in the context of balancing social needs with environmental
conservation.
STATE OF THE ECOSYSTEM
The status and trends of the environment and organisms in
the marine pelagic ecosystem, focusing on the upper 200
m, were recently evaluated in detail (Verity et al. 2002).
Salient conclusions from that analysis are briey sum-
marized below as a preface to recommendations for the
future.
The harvesting of at least 90100 million tonnes of sh
per year globally, and perhaps as high as 130190 million
tonnes per year (including by-catch), is the largest
anthropogenic impact on aquatic biota. Fisheries are
switching to younger sh, increasing their by-catch, and
targeting smaller species lower on the food web. Coupled
with long-term declines in those very stocks of sh of low
trophic level, more and more regions are likely to experi-
ence sheries collapses in the future (Pauly & MacLean
2003; Chapters 1 and 19).
At the ocean margins, an epidemic of nuisance
phytoplankton blooms is occurring, accompanied by
marine mammal, sh and invertebrate die-offs, human
deaths and illness, and signicant nancial loss to aqua-
cultural, sheries and tourism industries, causing food-
web dysfunction. Once considered to be rogue blooms,
they are now commonplace. Regions previously free from
harmful algal blooms now suffer such events, species
previously benign have become toxic or nuisances and, in
many regions, the frequency and intensity of red-tide
outbreaks have been increasing (Smayda 1997; Steidinger
et al. 2004).
Bioinvasions of alien species (plankton, macroalgae,
benthic invertebrates and vertebrates) are now common-
place (Ruiz & Carleton 2003), and often the result of rapid
and repetitive transoceanic or regional dispersal via
transit in ballast water or by shellsh transplantation.
Ecosystem invasions by non-indigenous jellysh, medu-
sae and ctenophores are increasingly commonplace.
These takeovers by exotic gelatinous organisms are
often associated with eutrophication and/or uncontrolled
shing (Purcell et al. 1999; Mills 2001).
It is becoming evident that the number of severely
depleted species of marine mammals, turtles and sh is
unprecedented in human history. Many marine species,
including some sharks and tunas, are considered endan-
gered (see Hilton-Taylor 2000), and many are likely to be
rendered extinct (Roberts & Hawkins 1999; Ellis 2003;
Chapter 1).
Chronic illnesses, disease epidemics, morbidity and
mass mortality events are being observed across an array of
taxonomic groups. A variety of pathogens, invasions of
alien and often toxic species and human illnesses appear to
be increasing in frequency and spatial extent (Harvell et al.
1999). These are provocative indicators at the ocean mar-
gins of a decline in marine pelagic ecosystem health (see
Verity et al. 2002 for a thorough treatment).
These vectors often interact in ways that are not easily
predicted and difcult to eliminate once established. They
signal change, degradation and ongoing cryptic reorgan-
ization of marine communities, leading towards a new
equilibrium driven by the need to adapt to multiple marine
ecological disturbances.
312 P. G. VERI TY ET AL.
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SUSTAINABILITY OF HUMAN INFLU-
ENCE: AN EXAMPLE
It is particularly inappropriate to put sheries in the general
category of natural-resource management along with for-
estry, not only because of the timescale problem already
discussed, but also because the sea is considered as the
common heritage of mankind with a lack of any ownership
for the sea, the seabed and the sh (Steele & Hoagland
2003). Unlike those on land, the resources and physical
characteristics of the ocean and seabed are typically con-
sidered to be either res communis, belonging to the public, or
res nullius, belonging to no one. In either situation, the
absence of private rights to a scarce resource, such as sh,
often leads to the tragedy of the commons (Hardin 1968), a
concept that arose not from forestry but from farming.
During the seventeenth and eighteenth centuries, the
establishment of long-term property rights in land, and their
allocation by owners to tenant farmers, became the sus-
tainable solution to an overexploitation of the commons.
The comparison of sheries management with farming
suggests two options that can be considered generically as
American and European (Steele & Hoagland 2003). In
the former, market forces have led to very large farms often
owned by even larger corporations who necessarily take a
long-term view. Such farms are, typically, very dull aes-
thetically but nancially efcient. The European alter-
native is to maintain a geographically and culturally
traditional landscape. This is aesthetically pleasing, but
requires central support by subsidy and import controls.
Each approach is a solution for a sector of the economy
which, although signicant politically, is not critical for
highly industrialized countries.
The developed world now depends more and more on
developing countries for sh supply so that the mature
sheries at high latitudes are even less critical economically
but are still culturally visible. For example, Europe and
North America try both to require economic efciency and
expect cultural continuity (Kurlansky 1997). It is not
surprising this leads to failure. Attempts to implement
programmes of marketable property rights in sh, through
such measures as individual tradable quotas (ITQs), have
been subject to moratoria in countries like the USA (NRC
[National Research Council] 1999). Alternatives involving
effort limitation are recognized to aggregate or integrate
shing effort into larger management blocks that neces-
sarily take a longer-term view but may deny independence
to individual shers; they are also likely to decrease the
number of boats and people employed. At present the
shers, rather than the sh, are regarded as endangered. As
the costs of enforcement of present policies increase and
the sh stocks decline, it will be interesting to see whether
free-market forces begin to trump the proponents of cul-
tural preservation. Certainly in countries such as Iceland or
New Zealand, where the shing economy is critical to
national economic production, institutional changes in the
direction of property rights and market allocations have
been seen (NRC 1999).
The European agricultural solution often results in
part-time farmers. This is not feasible in open-sea shing
given the nature of the work and the substantial capital
depreciation. It may be successful in nearshore sheries for
high-priced shellsh. The logical outcome for all these
problems seems to be sh-farming. In northern Europe,
the rearing of salmonids has converted an expensive deli-
cacy into a supermarket commodity comparable to chicken.
It may seem that, when the open sea has become bereft of
directly edible sh, it will then supply an enhanced source
of protein for the sh farms, from small rapidly growing
sh further down the food web (Chapter 1). This form of
enclosure is an obvious consequence of the inability to
evolve a management protocol for the open sea. Ironically,
such a combination, namely harvesting at lower trophic
levels to feed higher-level farmed stocks, could still lead to
greater natural exibility in marine ecosystems, such as
having an abundance of jellies rather than small sh at the
top of the food web (Chapter 1).
The present focus on sustainability, especially of indi-
vidual sh stocks, ignores the natural variability of marine
ecosystems at decadal scales (Steele & Hoagland 2003).
Attempting to damp out these interannual trends by year-
to-year management practices may even exacerbate the
oscillations, particularly by compounding them with eco-
nomic processes at the same timescales. The natural
physical and ecological causes of these cycles need to be
understood and sufciently long-term management then
revised to ameliorate rather than amplify the economic
consequences.
NATURAL OSCI LLATIONS
AND ANTHROPOGENIC IMPACTS
TO THE YEAR 2025
For many purposes, the ocean could once be regarded as a
vast reservoir relative to human activities; anthropogenic
impacts could usually be assumed to be restricted to local
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areas. With the combination of human population increase,
changes in consumption patterns and technological changes
in activities such as shing, obvious impacts have spread
into large offshore regions. For example, the collapse of
Atlantic cod sheries (Hutchings 2000) and anoxia events in
North Sea bottom waters (Karlson et al. 2002) are warnings
of future trends in the absence of appropriate actions.
Human activities have a direct impact at both ends of the
oceans pelagic food chain. At the upper end of the food chain,
the impact of large-scale sheries and hunting on marine
vertebrates (mammals, reptiles and birds) potentially cascades
down through the food web, creating indirect inuences at
lower trophic levels (Verity & Smetacek 1996). Likewise,
changes in nutrient load and nutrient stoichiometry on the
lowest trophic levels potentially cascade upwards through the
food chain to create changes in rates, biomasses and bio-
diversity at the high end. This pelagic food chain contains life
forms from viruses to whales, covering eight orders of mag-
nitude in size, forming a dynamic system where important
processes cover timescales from fractions of a second to
decades. Even if good conceptual models existed to cover
these ranges, the net effect of forces working from opposite
ends of the food web would probably be hard to predict.
Despite such difculties, there is a relatively well-
accepted paradigm for the qualitative changes expected in
the structure of the lower part of the food web along natural
gradients from oligotrophy to eutrophy. In oligotrophic
waters, primary production is dominated by picoplanktonic
(size <2 lm) organisms, while larger-celled algae dominate
in more eutrophic conditions. Combining the phytoplank-
ton with their microbial predators (protozoans) and with the
degraders of organic matter (heterotrophic bacteria), this can
be summarized in a kind of ladder-like food-web structure.
Such a picture is an elaborated view of the apparent coun-
ter-intuitive observation that nutrient-rich environments
are dominated by short food chains, such as the classical
diatommesozooplanktonsh pathway (Fig. 20.1), while
nutrient-poor environments are dominated by long
food chains (e.g. autotrophic picoplanktonheterotrophic
agellatesciliatescopepodssh) (Ryther 1969).
Even from highly simplied views of the pelagic food
web (Fig. 20.1), it is obvious that human activities have an
impact at the bottom of the food web. Not only will the
amount of dissolved nutrients released to the environment
around the ocean margins have an impact through stimu-
lation of microbial growth, but the qualitative composition
of such releases to the ocean is also important. Organic
pollutants, even when non-toxic, change the environmental
forcing by fuelling substrate into the heterotrophic bacteria
(Fig. 20.1), potentially shifting the ow of matter, energy
and nutrients toward long food chains from bacteria via
heterotrophic agellates and ciliates to copepods, before
Heterotrophic
flagellates
Ciliates Mesozooplankton
Approximate size (m)
Diatoms
Autotrophic
flagellates
Autotrophic
picoplankton
Heterotrophic
bacteria
Dissolved
organic matter
10
7
10
6
10
5
10
4
10
3
10
2
CO
2
Fish
Jellyfish
Fig. 20.1. Idealized representation of carbon ow through the pelagic food web, indicating how organic carbon may follow a long
pathway, starting from small autotrophic picoplankton, or a short classical pathway, starting with diatoms before passing through
mesozooplankton to enter the higher parts of the food web ending in either sh or jellysh.
314 P. G. VERI TY ET AL.
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reaching trophic levels of potential commercial interest.
Dam-building in rivers has the effect of reducing silicate
transport to the ocean (Conley et al. 2000). Silicate limi-
tation will restrict the potential for establishment of the
short food chain from silicate-dependent diatoms via
copepods to sh. Dam-building in the Danube River
appears to have had such an effect on the Black Sea
ecosystem (Humborg et al. 1997).
Modern agricultural practices in industrialized countries
tend to release large amounts of nitrate relative to phosphate
in runoff waters, whereas releases of untreated domestic
sewage are relatively enriched in phosphate. In places like
northern Europe, the consequence is a linking of coastal
ocean chemistry to meteorological conditions: winters with
high precipitation and large water ow in the rivers lead to
excess transport of nitrate to the North Sea (Hickel et al.
1993), with phosphate and silicate-limited growth of the
primary producers as the expected consequence. Harmful
algal species often exhibit diverse nutritional strategies
including direct consumption of organic nutrients as well
as predation on competing phytoplankton. Hence, coastal
zone management strategies may not only change ecosystem
functionality, but also the stoichiometric relationship
between elements and the way the elements are tied up in
inorganic and organic forms, with potentially severe eco-
system consequences (Anderson et al. 2002a).
Aeolian inputs to the ocean may contain substantial
inputs of nitrogen, phosphate and iron. In the Mediter-
ranean Sea, northerly winds may carry nitrogen-enriched
aeolian inputs characteristic of the industrialized areas of
Europe, while winds from the south carry Saharan dust
enriched in phosphate and iron (Bethoux et al. 1998;
Herut et al. 1999). Long-term changes in dominating
wind patterns, for example by extended changes in the
pressure difference between the Azores and Iceland, may
thus inuence the biogeochemistry of the Mediterranean
basin. Adding to the complexity of climate change effects
on the pelagic ecosystem is the interaction between dust-
transported iron availability and the high iron require-
ment of the nitrogen-xing enzymes. The iron in Saharan
dust transported into the Atlantic has thus been suggested
as a mechanism stimulating nitrogen xation in the
Sargasso Sea ecosystem to a degree that may shift this
system to phosphate limitation (Wu et al. 2000).
One of the predictions of climate models, in addition to
a temperature increase, is increased instability of weather
patterns. Upwelling events where nutrients are brought up
into the photic zone are frequently wind-driven. Typically,
such events are believed to stimulate the classical food
chain of diatomsmesozooplanktonsh (Fig. 20.1),
potentially enhancing sh production, but also being a
short-lived phenomenon lasting only until most of the
imported nutrients are re-exported via mechanisms such as
sinking diatoms and copepod faecal pellet production
(Wassmann 1998; Boyd & Newton 1999). In such a scen-
ario, low-pressure passages in resonance with the food-web
transfer of energy and matter through the classical part of
the food chain could be very benecial for sh production.
Mixing events that were insufciently frequent would
push the system towards long pathways (Fig. 20.1), while
too-frequent events would not allow the upwelled nutri-
ents to be translated into biological production in the
photic zone (Boje & Tomczak 1978; Payne et al. 1992).
Socioeconomic decisions may thus inuence the oceanic
environment, not only through land use and wastewater
treatment, but also via the longer loop, where greenhouse
gas emissions potentially change wind and/or precipitation
patterns and thus nutrient load and stoichiometry in coastal
seas. In the North Pacic Central Gyre, a decrease has
occurred in silicate and phosphate over the last three dec-
ades (Karl et al. 2001). This has been suggested to be
caused by long-term, climate-driven changes that have
induced a domain shift in the microbial food web towards
dominance of small prokaryotes (dominance of the left side
of Fig. 20.1).
Also spectacular is the regime shift higher up in the
food web in the Black Sea, where a large part of the pro-
duction has been shifted into the commercial dead-end of
jellysh biomass (Berdnikov et al. 1999). As opposed to sh
larvae that seek out their prey by visual means, jellysh are
non-visual predators. One hypothesis for such shifts has
thus been that decreased water clarity (e.g. caused by
humic substances or increased content of inorganic or
biogenic particles) may induce such a regime shift towards
jellysh dominance (Eiane et al. 1999). However, under-
standing of the driving forces, and knowledge of when to
expect such shifts, is poor.
The collapse of populations at the high end of the food
chain, such as the North Sea herring, the Grand Banks
(USA) cod stock (Hutchings 2000) or the large whales
(Roman & Palumbi 2003) are well known (Chapters 1 and
19). The potential cascading effects down to the lower levels
in the food web remain, however, mostly speculative. Part of
the reason for this is perhaps the dichotomy suggested by
the simplied food web (Fig. 20.1). In the world of uni-
cellular organisms at trophic levels lower than those of most
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copepods, characteristic timescales for most processes are
those of days or less. In many cases this is short relative to
the hydrographical mixing processes of the water column, as
demonstrated by the fact that stable vertical structures such
as deep chlorophyll maxima become established (Steele
1978, 1995). At the level of copepods, the typical timescales
for reproduction increase to weeks or months, while the
timescale of vertical motion may be of hours to days. The
result is that the coupled processes of copepod feeding and
growth in the microbial food web may be separated in time
and space (Hansen et al. 1997). These differences in char-
acteristic timescales between trophic levels increase again to
the next predator levels, with the result that coupled pro-
cesses become much more difcult to demonstrate and
model above the microbial level.
WHERE NEXT: GLOBAL SCIENCE
OR LOCAL I SSUES?
Science appears to have come to the end of the rst
generation of global ocean programmes (World Ocean
Circulation Experiment [NODC 2003], Joint Global Ocean
Flux Study [JGOFS 2008] and Global Ocean Ecosystem
Dynamics [GLOBEC 2004]). Although the original
impetus, two decades ago, for a coherent suite of pro-
grammes came from the USAs National Science Founda-
tion, these programmes became international in content and
participation. Their development increased the scale of the
science, required greater integration of management and
became dominant themes in marine pelagic research (NRC
2000b). But there is now increasing emphasis on the
application of the science to societal issues, including pol-
lution, climatic change and sheries (Field et al. 2002).
These require better integration across the sectors. As past
work is built on, but these growing concerns are also
responded to, are global programmes with their sectoral foci
the best way forward? Or, as has been suggested (NRC
2000b), should the emphasis shift to a loose coupling of
regional or local problem-based programmes?
Current studies of the functioning of pelagic marine
systems are based largely on modelling the responses of
particular populations to physical forcing. These studies
have been very successful in demonstrating the rst-order
role of physical processes in determining the distribution
and abundance of copepods, krill and sh larvae (Steele &
Collie 2005). However, the biological controls within
marine ecosystems that act through density-dependent
feedbacks across trophic levels need to be understood.
These processes are essential for the longer-term persist-
ence of systems and especially their ability to switch among
different, but ecologically coherent, regimes (Verity &
Smetacek 1996; Steele & Collie 2005). The comparison
of ecosystems in contrasting locations should not only
demonstrate the different physical controls, but can also
help elucidate possible underlying ecological similarities or
constraints. For this, data and concepts from all trophic
levels need to be incorporated.
One outstanding feature of past and current global
projects has been the lack of real interaction or overlap
between studies of the upper and lower parts of marine
food webs. This may have been because: (1) the research
interests vary greatly among groups (e.g. global carbon
cycle versus sheries studies); (2) the regions studied do
not usually overlap (e.g. the open ocean versus continental
shelf); (3) the methodologies are distinct (e.g. bio-
geochemistry versus taxonomy and population dynamics);
and (4) the research groups are often separated by location
and funding sources.
However, research as well as social concerns have
brought these groups closer together. For sheries,
addressing the vexing questions of causality of sh popu-
lation declines, namely overshing (topdown effects)
versus environmental change (bottomup and topdown
effects) (Chapter 1), requires much more attention to lower
trophic levels than has been traditional, as well as more
integrative studies combining ecological and food-resource
approaches. Similarly, questions about the ecological
consequences of longer-term climate variability require
more study of the effects of changes in carbon and nutrient
export on recycling through mesopelagic and benthic food
webs. These issues make it imperative that future projects
consider marine food webs in their entirety, from phyto-
plankton and other microscopic organisms all the way to
top carnivores.
A key aspect is the export of organic matter into the
oceans interior. The fraction of total primary productivity
(PP) that is exported to deeper water or to higher trophic
levels (including sh) depends on, but is not directly
related to, the rate at which new nutrients are introduced
to the euphotic zone (Iverson 1990; Steele & Collie 2005).
This new production (or the corresponding f-ratio new
production/total production) is difcult to determine
experimentally. Many estimates of transfer to higher
trophic levels or export to deeper water are extrapolations
from a few measurements or rely on estimates from models
(e.g. Pauly & Christiansen 1995). Also, the denition of
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new production depends on the relevant timescale as well
as on the physical structure of the water column. Nearly all
the recent f-ratio determinations (e.g. JGOFS) have been
for open-ocean waters and should not be extrapolated to
coastal regions (e.g. Chapter 19), for which there are few
data. This is particularly true for regions subject to heavy
sheries exploitation, where the main concern is the
availability of new production to higher trophic levels
(Steele & Collie 2005). If management is to become eco-
system-based, good estimates will be needed of the portion
of primary production ultimately available to higher
trophic levels, principally sh, but also marine mammals,
sea turtles and seabirds.
Great cycles occur in the natural abundances and
geographic locations of sardines and anchovy, with periods
of a few decades (Worm & Myers 2003). Dramatic switches
in the abundance of major ground sh stocks have been
witnessed on Georges Bank off New England (USA),
where cod were replaced by dogsh and mackerel, while
off Newfoundland crustaceans became abundant after
the cod went (Chavez et al. 2003). These switches can
have great economic consequences, especially on shing
communities. Yet, these large rearrangements in species
composition appear to be ecologically acceptable responses,
using alternative pathways within the higher trophic levels
of the ecosystem. There is little evidence of any signicant
effects on primary productivity.
Climate change may increase ocean temperature and
stratication, thus reducing new production by nutrient
input from depth. Together, these physical trends could
create signicant non-linear alterations in export to both
the deep ocean (Chapter 22) and upper trophic levels.
Thus, changes in production do not translate simply into
corresponding changes in yields of organisms at higher
trophic levels, for example pelagic and demersal sh.
Conversely, there are issues concerning the consequences
of the dramatic changes in abundance of sh stocks to
other food-web components. Key questions include: do
these changes have impacts on the microbial components
in food webs through release from topdown controls? Or
do transports from the large zooplankton (copepods) and
by detritus act to partially decouple feedback between
upper and lower food-web levels? Or is this distinction a
result of the dichotomies in the way the research is
structured (Verity et al. 2002)?
Physical forcing on marine systems is related to winds,
circulation, density fronts, gyres and tides. All these forcing
factors are associated with very different time and space
scales, and their biological consequences can occur at a
considerable distance in time and in space from the site
where the energy input occurs. Further, responses of marine
food-web components to these factors are not necessarily
linear. This has led to the concept of regime shifts
(Beamish 1993), in which gradual long-term changes in
ocean physics can lead to relatively rapid switches from
one community structure to another. This has major
implications not only for the potential to forecast sh stock
changes, but also for attempts to infer climatic changes from
palaeo-oceanographic records. Rigorous testing of the
underlying hypothesis of multiple equilibrium states (Steele
& Henderson 1984; Collie & DeLong 1999) requires much
more information on food-web dynamics, especially at
intermediate trophic levels.
The consequences of physical forcing in terms of sec-
ondary production may differ greatly between coastal and
open-ocean areas. In coastal waters, the mixed layer can
extend to the bottom, affecting benthic processes (see
Chapter 19). In the open ocean, the mixed layer is shallow
with respect to the sea oor, so processes that occur in the
mixed layer will inuence the benthos less immediately
than they do inshore.
Heavy shing tends to remove the larger, more com-
mercially valuable sh, leaving primarily the smaller, less-
valuable sh (Pauly et al. 1998; Myers & Worm 2003).
There is observational and theoretical evidence that such
large changes at the top of marine food webs can induce
switches in equilibrium states at lower trophic levels
(Spencer & Collie 1997). Such effects can lead to top
down regime shifts in which the system sustains a different
community of sh and other species. Such an alternate
state can be stable and may need a strong perturbation to
shift it back to its previous, or other, state (Collie & De
Long 1999). Historically, these switches appear to occur on
decadal to centennial timescales but, under strong forcing,
could become more frequent (Steele & Henderson 1984).
CONCLUSI ONS
Scientic perspective
All of these issues come together in the context of human
impacts on the diversity of marine life (Steele & Collie
2005). How can human use of marine resources and the
maintenance of adequate species richness to sustain marine
ecosystem structure and function be balanced? In sheries,
for example, three apparently simple questions need to be
answered. (1) How diverse are the resources available to
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humans? (2) How much can be removed by humans?
(3) How sustainable is the yield? Answers to such questions
require an understanding of the statics (food-web structure),
the kinematics (energy and nutrient uxes) and the
dynamics (system stability) of ecosystems. Yet, effects of
differences in marine biodiversity on marine food-web
kinematics remain unknown (Grassle 2000). The main
changes observed in marine species are at the larger end
of the size spectrum in the macrozooplankton and sh.
Yet most of the marine biodiversity lies at the other end of
the size spectrum: in the viruses, bacteria, algae and
microzooplankton of the microbial loop. Very little is
still understood about how external factors inuence
biodiversity.
Interest in longer timescales has consequences for the
spatial scales of study. In the ocean, both physical and
ecological scales of time and space are inextricably linked
(Steele 1995). The processes of mixing and transport of
water masses change with increasing time and space scales;
studying biological processes on longer temporal scales
implies the need to study processes at larger spatial scales.
Thus, scales of interest now have to extend from the
micro- and mesoscale to basin and global scales
(Schwartzlose et al. 1999). This increasing spatial range
imposes greater demands on theory and eld programmes.
In particular, the longest temporal scales require retro-
spective data, such as those obtained from sh scales
(Baumgartner et al. 1992).
The focus on physicalbiological coupling at different
trophic levels in JGOFS and GLOBEC has brought sci-
entic benets. At the same time, the development
of linearized food-web models (Vezina & Platt 1988;
Christensen & Pauly 1993) can provide comprehensive
pictures of trophic structure by ignoring the non-linearities
imposed by the physics and community dynamics. Yet,
there is increasing evidence that these non-linearities may
be expressed by abrupt or discontinuous changes in com-
munity structure, namely regime shifts (Steele 2004). How
pervasive such hysteresis loops are in natural systems
remains to be seen (Scheffer et al. 2001). But they can have
severe consequences for management, as witnessed by the
lack of revival of cod stocks in the north-west Atlantic.
Thus a precautionary approach would assume that such
alternative stable states would occur in marine ecosystems
under stress; once changed, ecosystems may stay that way.
Philosophical perspective
The secular impacts of humans on the oceans represent a
staggering diversity of vectors and accumulated injustices
(Verity et al. 2002). There can be little doubt that, with
burgeoning human population, continued evolution of
human society on a global scale is predicated on mutually
benecial moral ethics and altruistic behaviour. There
exists a continuum of perspectives on how humanity
should balance social needs with those of environmental
conservation (Gorke 2003).
At one end of the spectrum, the fundamental conser-
vationist view is that the pelagic ecosystem is crying out for
help from the activities and by-products of a species that,
for the rst time in the history of the planet, is capable of
altering evolution. Conservation theory simply holds that
humanity must look beyond immediate resource use and
long-term use of the oceans as a receptacle for human
organic and technological by-products.
At the opposite end of the continuum are similarly
compelling arguments couched in social consciousness. To
wit: in a world with an exponentially growing population,
the immoral thing might be to not actively pursue use of
the oceans and their resources for humanitys benet. The
overriding constraint from both a moral and a practical
point of view, of course, is how to ensure sustainability.
The only long-term solution to this Malthusian prob-
lem in the oceans is to achieve control of the global human
population explosion. Everything else is a variation on the
famous shell game, moving the problem around, perhaps
delaying things a decade or two, but without long-term
solution. Ideally restoring the marine pelagic ecosystem to
its state when the rst shing people baited hooks, to the
time when human ancestors looked for wealth on land but
were unable to recognize it in the oceans, might be dreamt
of. This is undoubtedly impossible in the present global
economy and political climate. But if humans are not aware
of the breadth and depth of the problems, then society is
not prepared to plan for effective and intelligent policy and
action. The warning signs in the past have largely been
ignored to date: evidence suggests that continued ignor-
ance will lead to changes in the structure and function of
the marine pelagic ecosystem that will overshadow every
future human step taken, and that is likely to produce a
new equilibrium that will be less anthroponomic.
318 P. G. VERI TY ET AL.
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