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The use of ecosystem characteristics in fisheries

management
a
Spencer Apollonio
a
31 Eastern Avenue, Boothbay Harbor, ME, 04538
Published online: 23 Dec 2008.

To cite this article: Spencer Apollonio (1994) The use of ecosystem characteristics in fisheries management, Reviews in
Fisheries Science, 2:2, 157-180, DOI: 10.1080/10641269409388556

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 Reviews in Fisheries Science, 2(2): 157-180 (1994)

The Use of Ecosystem Characteristics

in Fisheries Management*
Spencer Apollonio
31 Eastern Avenue, Boothbay Harbor, ME 04538

* This is a modified version of an article originally presented by request at the National Ecosystems
Conference, Orlando, FL, March 22-23, 1988, organized by the National Marine Fisheries Service, National
Oceanic and Atmospheric Administration.
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ABSTRACT: Application of the concept of ecosystems to fisheries management would

provide managers with a useful conceptual framework within which they may (1) absorb
specific scientific advice, (2) anticipate probable resource responses to fishing perturbations,
(3) anticipate species interactions, and (4) anticipate the effects of management measures.
Ecosystems in general encompass three major components: (1) species life history strategies,
(2) the evolution of communities, and (3) hierarchic, rate-regulating structures. The interrelation
of these components and apparent examples in fisheries ecosystems are reviewed. Most
fisheries experience a reversal of the natural evolution of ecosystems dynamics, that is, fishing
induces greater frequency and amplitude of variations of specific components of the system.
An understanding by managers of the functions and interrelations of the basic components
of systems clarifies the reasons for the typical response of fisheries ecosystems to the fishing
effort. The implications of ecosystems characteristics for fisheries managers are also briefly
considered.

KEY WORDS: ecosystems fisheries management, fisheries ecosystems, hierarchies in fisheries.

/. INTRODUCTION
Fisheries managers are routinely inundated with data about a multitude of species.
Usually they consider these data with reference to only one species at a time,
although they are aware that their responsibility is for a community of species, not
just a collection. Managers are almost forced to consider only one species at a time
because they frequently lack of community conceptual framework within which to
absorb, evaluate, and act on the plethora of species information.
The information coming to managers from research efforts could be cast into a
useful conceptual outline or framework of fishery ecosystems. That is, managers
should have a mental image of their system in simple, general terms that can be
related to the diversity of scientific information they receive that would permit them
to make some general estimates of the consequences of fishing and management
actions on the system. Any community of fish species is part ôf a larger marine
ecosystem, and the missing conceptual framework may be how ecosystems, in
general, are structured and how they work.
1064-1262/94/$.50
©1994 by CRC Press 157
 APOLLONIO REVIEWS IN FISHERIES SCIENCE

An understanding of the characteristics of ecosystems would provide managers

with a powerful asset. It could explain the inherent and characteristic variabilities
of various species as well as the influence of environmental factors and fishing effort
on those variations. It also could provide a predictive capability for the probable
trends of abundance of fish species, or even, whether such predictions are possible
for certain species. A knowledge of which species are amenable to forecasts of
trends in abundance is important for managers.
With a sufficient understanding of ecosystems, one should be able to determine
those species that are legitimate candidates for efforts at stock stabilization and those
that are inherently highly variable. Current management practice appears to promise
a sustained yield for all species, independent of their biologic peculiarities, if only
the proper regulations were promulgated. It appears as though managers are now
almost required to consider that all species are biologically equivalent or that
characteristics of different species are without management significance.
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Managers need three basic kinds of information:

1. The consequences of various levels of fishing effort on the stocks and on
long-term yield potential
2. The consequences of proposed management measures
3. The fundamental or conceptual reasons for each of those consequences
within the functioning of ecosystems
In many instances, simple ecosystematic statements on these points would be useful
to managers and, in some instances, perhaps sufficient. However, fish managers, in
general (and, occasionally, fish scientists) do not have a common definition of the
term ecosystem. Until such a definition is available, the term can hardly be a unifying
concept or testable hypothesis for directing research or management. As long as the
term remains undefined, vague or anecdotal, it will remain a concept of marginal
utility.
Science advances through the process of testing refutable hypotheses. Fisheries
science is periodically criticized because of a dearth of refutable hypotheses
(Walford, 1958; Hennemuth, 1979; Laurence Ms, 1986; Rothschild and Osborn,
1986). Before the ecosystem concept can serve fisheries management, it must be
expressed in a more rigorous form. Unless such expression occurs, the ecosystems
approach may only burden managers by adding one more concept to many
approaches already in fisheries management. If the ecosystems concept were to be
formulated as testable hypotheses, it might redeem its promise as a productive basis
for management.

//. ECOSYSTEMS CONCEPTS

First, we should not permit definition of the term ecosystem to be anything that we
find to be convenient. If the term has no commonly understood, well-defined limits,
then it has no meaning and so has no use.
Second, although we may postpone a formal definition of ecosystem, we may
note, as Lloyd Dickie (1973) pointed out, that ecosystems have certain general

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 Volume 2 (Issue #2), 1994 ECOSYSTEM CHARACTERISTICS IN FISHERIES MANAGEMENT

properties. He addressed both of these issues — those of definition and those of

general properties — as follows:

1. Systems have a hierarchic organization.

2. Although there is interdependence in the hierarchy, among the "lower" or
less well-organized parts of the system there are relatively large variations.
Despite this, the overall function is sustained by the presence of homeostatic
mechanisms. The result is that in a system the overall variance is much less
than the sum of the variances of its parts. This is often the essence of the
meaning of the term system.
3. A system is not a machine because quite large perturbations of a part will
be compensated for in other parts, thereby preserving the functioning of the
whole. The overall function depends on the interrelations among parts but
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is more or less independent of the nature of the individual parts.

By recognizing the properties of systems as noted by Dickie, we may determine

whether we are studying, in fact, a real system or simply defining a convenient, but
arbitrary, aggregation that may have no meaning within the systematic functioning
of the ocean. If we are to manage resources within an ecosystem context, then we
must attempt to manage with reference to real systems, not arbitrary and convenient,
but catchall assemblages.
Ramon Margalef (1975) has offered a perspective on ecosystems that supple-
ments and complements Dickie's. Margalef identifies three major components
essential to the definition of ecosystems:

1. Single-species life history strategies

2. The evolution of life history strategies, or, in fact, the evolution of the nature
of communities of species
3. The mechanisms or linkages among species that regulate the systematic
functioning (in Dickie's sense) of the community.

Margalef stated that "... the strategy of single species during evolution, their
replacement in the course of succession, and the action of many natural mechanisms
that slow down the turnover in ecosystems, can be encompassed in a general view
of ecosystem dynamics." — a summary of the qualities of an ecosystem that is as
succinct as any we could probably find.
From many pieces of evidence, the general structural and functional features
outlined by Margalef may be accepted as ecosystems realities, and, in fact, may
provide components essential to a formal definition of a fisheries ecosystem. The
interactions of these components form the basis and are the mechanisms responsible
for the ecosystem characteristics and their emergent properties as identified by
Dickie.
The three components proposed by Margalef may be restated in the form of
hypotheses, which have been explored at length and are broadly accepted by
ecosystems ecologists. Each hypothesis will be examined in turn.

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 AP0LL0N10 REVIEWS IN FISHERIES SCIENCE

///. THE HYPOTHESIS OF LIFE HISTORY STRATEGIES

The first hypothesis, in a sense, states the obvious, that is, species differ in
fundamentally important ways. Further, and more interestingly, the differences
appear to form two generally distinct categories of species but that actually form
continua from one kind to the other. These kinds are called r-selected znd K-selected
species, and their general attributes were proposed by MacArthur and Wilson (1967)
and summarized by Pianka (1970), Gadgil and Bossert (1970), Stearns (1976), and
others. Whereas Pianka listed "climate" in his original table of correlates Cable 1),
undoubtedly "environment" in the broad sense "from the species' point of view"
(Begon and Mortimer 1981), or even "niche" (Root 1967) conveys a better sense of
the meaning of the correlates.
A cautionary note may be appropriate here. Although there is criticism of the
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validity of the r-^Tconcept, partly because it fails to account universally for all aspects

TABLE 1
Some of the correlates of r- and /(-selection
Correlates r-Selection
Climate Variable and/or unpredict-
able, uncertain
Mortality Often catastrophic, nondirected,
density-independent
Survivorship Often type III (Deevey,
1947)
Population size Variable in time, nonequilibrium
usually well below
carrying capacity of
environment; unsaturated
communities or portions
thereof; écologie vacuums;
recolonization each year
Intra- and interspecific
competition Variable, often lax
Relative abundance Often does not fit MacArthur's
broken stick model (King,
1964)
Selection favors Rapid development
Early reproduction
Small body size
Semelparity: single reproduction Larger body size
Length of life Short, usually less than 1
year
Leads to Productivity
From Pianka, E. R. Am. Nat., 104:592-597 (1970). With permission.
K-Selection
Fairly constant and/or pre-
dictable, more certain
More directed,
density dependent
Usually type I and II
(Deevey, 1947)
Fairly constant in time, equi-
librium; at or near carrying
capacity of the
environment; saturated
communities; no recolorization
necessary
Usually keen
Frequently fits the MacArthur
model (King,
1964)
Slower development,
greater competitive ability
Lower resource threshold
Delayed reproduction
Iteroparity: repeated productions
Longer, usually more than
1 year
Efficiency

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of life history strategies, nevertheless, it is a useful paradigm of the reality of

qualitatively different kinds of strategies. Begon and Mortimer (1981) stated that the
concept "... has been a notable success in the ecologists' search for pattern.... It
may indeed be the single most useful simple classification..." This is the kind of
qualitative ecosystem generalization that has potential value to managers.
It must be understood that species are not r- or AT-selected in absolute terms,
fixed immutably only at the extremes of the scale. Various species, in fact, form
continua, each continuum appropriate for a system, grading progressively from one
life strategy to the other. Individual species may be at various points of a continuum,
each relatively more or less r- or Jf-selected with respect to others.
The terms r- and K- are, of course, simply a form of widely recognized
shorthand, symbolizing not just one but a group of life history characteristics, which,
acting as a whole, determines the life history strategy of the species. Briefly, the
terms r and K are derived from parameters of the growth (or logistic) curve for a
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population (Figure 1), r symbolizing species that maximize reproductive potential,

and K symbolizing species that have evolved toward the most efficient use of
resources and thereby maintain a population level close to the carrying capacity of
their habitat. Perhaps more descriptive terms for these species are opportunistic for
r-species and equilibrium or competitor efficient for ^-species. These terms convey
a descriptive sense of the functions of the species within the ecosystem and
summarize the fundamental species characteristics of interest to managers.
Is a continuum of species useful to managers in any practical sense, or is it
simply an abstract, conceptual blur? This question appears to be a principal criticism
of the hypothesis. In fact, it appears that species may be quantitatively related to
each other on the continuum, as explicitly demonstrated by Gunderson and Dygert
(1988) (Figure 2) and implicitly by Hennemuth and Shepherd (1983) (Figure 3), and
therefore, these attributes are helpful to managers. May (1978) and Adams (1980)
urged that any rational conservation or management plan should begin with
consideration of this hypothesis of r- and ^T-selection, which has rarely, if ever, been
considered in marine fishery management (see Kawasaki, 1980).
This neglect is surprising because the hypothesis indicates to managers that
some species (r-selected) are inherently more variable and, therefore, probably

TIME

FIGURE 1 . The concepts of r and K with reference to a generalized curve of population growth.

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0.0 0.1 0.2 03 0.4 0.5 0.6 0.7

GONAD INDEX (WG3)
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FIGURE 2. Characterization of instantaneous natural mortality [M] for 20 stocks of fish derived from
wet gonadosomatic weight indices. (From Gunderson, D. R. and Dygert, P. H., J. Cons., 44:200-209
(1988). With permission.)

11
10
CH PHYTOPLANKTON
LU
ta
3
Z 10 ZOOPLANKTON
<
HI • HADDOCK
10 HERRING
3

WHALES

-1 3 5 7'
10 10
10 10
10 10
10 10

STRATUM VARIANCE

FIGURE 3. Statistical analysis of fisheries aggregations. (From Hennemuth, R. C. and Shepherd,

S. P.. ICES. C M . 1983.)

unmanageable in the usual sense and do not, in fact, particularly need management.
It states that other species (/T-selected) are manageable by traditional methods and,
more importantly, are particularly vulnerable to fishing effort without management
restraint. Adams (1980) made these points persuasively.
The concept explains why the basic Beverton Holt yield model works for some
species but not for others (Dickie, 1971; Ursin, 1984).

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The relative positions of species on the r-K continuum could be determined by

comparison of various parameters of their life histories. Gunderson (1980), basing
his argument on the concept, estimated natural mortality rates from gonadosomatic
indices and, to that extent, arranged species on a continuum. On the basis of at least
10 of 12 parameters, herring seems to be more JiT-selected that cod (Table 2),
although this table should be considered heuristic rather than definitive. It is the
aggregate of parameters, however, that constitutes a life history strategy. A general
formula probably could be developed that would properly weight and combine
significant parameters into a measure of a species position along a scale of life
history strategies.
There are at least two substantive reasons for ranking species on the r-K
spectrum. The first is that it could predict the degree to which a species is amenable
to management in the sense of Adams (1980), and it could also predict the possible
response of species to fishing effort (Figure 4). The second is that it could predict
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the relative position of the species within the hierarchy of the ecosystem, and, thus,
the extent to which it is a constraining or regulating force within the system, as a

TABLE 2
A tentative comparison of the relative
r- and /(-selection of cod and herring
Cod Herring
% Mature at age 3 75% 25%
Egg numbers 1,000,000 10,000
Fecundity/g female 475 300
Egg size (mm) 1.1-1.8 1.0-1.4
Egg habitat Pelagic Demersal
Larvae size (mm) 4.0 5.0-6.0
Yolk sac absorbed 4.5 10.0
Production/biomass 0.6 0.3
Food specificity Less More
Schooling behavior Less More
Aggregation Less More
Vulnerability to fishing Less More

10 10
GENERATIONS

FIGURE 4. Generalized responses of /(-selected (left) and r-selected (right) species to a single
perturbation (From Southwood et al., Am. Nat, 108(964):791-804 (1974). With permission.)

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 APOLLONIO REVIEWS IN FISHERIES SCIENCE

means of modulating fluctuations in the abundance of other species in the system.

There is a relationship between hierarchic structure and r- and /T-selected species
(Margalef, 1975). "Many strategies within an ecosystem appear to be complementary;
for instance, a predator may be a kind of ÜT-strategist (MacArthur and Wilson, 1967)
primarily adapted to biotic elements, while its prey may have to remain closer to the
r-strategy, geared primarily to abiotic factors." Further (Margalef, 1975), "... in the
evolution of species there is a higher probability of passing from r- to ^-strategy than
the reverse, and, in écologie succession, a stabilizing element for the ecosystems
consists of the increasing importance of species that establish connections more and
more extended over time and space. In practice, this amounts to K-strategist taking
over control of the ecosystevf (emphasis added). iC-species, because of their biologic
attributes, have a role in "structuring" or "shaping" their systems (explicitly Margalef,
1975; Estes, 1979; Ray, 1981; implicitly O'Neill et al., 1986). That is, their presence
induces predictability into the system, whereas, by inference, their reduction or
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removal reduces internal control mechanisms and increases variability and

unpredictability.
It is quite possible, for example, that the present character of the fisheries of
upwelling systems is a consequence of the removal of apex predators from those
systems. McCall (1986) estimated severe reductions of fur seals, sea otters, elephant
seals, sea lions, whales, and seabirds in the California current. Preexploitation
populations of these predators, by McCall's estimates, would have consumed
millions of tons of fish in the California system. These quantities would clearly
amount to substantial "structuring" of the system as well as significant biologic
impact, and, by the nature of the hierarchic structure of ecosystems, to greater
predictability within the system.
There has been the implication throughout the exploration of the r and K
concept in écologie literature that a species is fixed in its position on the continuum.
Terrestrial ecologists (Young and Muyshandt, 1972), however, suggested a shift on
the continuum for insects after disturbance of tropical forests by man. A shift for seals
and whales in Antarctica, as a probable consequence of severe reductions of blue
and fin whales, appears to be evident (Laws, 1985). In certain respects, it is obvious
that exploited fish species also show shifts (smaller average population size, fewer
spawning year classes, faster growth rates, earlier maturity, higher production-to-
biomass ratio) (Figure 5). In fact, the acknowledged conventional fishing strategy is
to shift the ratio of production-to-biomass to a higher level by restructuring the
unexploited population (usually through reduction of average age) to a point where
the optimum yield can be obtained by taking advantage of the more rapid growth
capabilities of younger fish. This strategy must inevitably affect other life history
parameters. One would expect, therefore, a population shift toward greater
r-characteristics. A consequence of this shift is greater uncertainty in population size
and increased amplitudes of variability (Figure 6). These characteristics have, in fact,
been reported in several heavily exploited fisheries (Steele, 1979; Hennemuth, 1979).
The shift of uncertainty and the increased amplitude of variability are both
predictable from the hypothesis. The probability that species shift along the r-K
continuum should be taken into account as an essential component of fisheries
science and as a predictable parameter with management implications. Moreover,
with respect to increased variability, Laurence (1986) noted that there are at least
three distinct definitions of the "recruitment problem" on which fisheries research

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10 - r— 1951
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2 3 4 5 6 7 8 9 1 0 10+
Age in years

FIGURE 5. The reduction of year classes of herring in the North Sea. Not shown is a concomitant
reduction of age at maturity. (From Cushing, D. H., Fisheries Biology, a Study in Population Dynamics,
University of Wisconsin Press, Madison. With permission.)

is increasingly focused. On the basis of these hypotheses and this shift, I suggest
another definition: the "recruitment problem" is that the qualitative and quantitative
natures of the forces controlling each year class strengths are changing in a manner
predictable from the initial character of the species or communities and also from
the magnitude of the fishing effort; that as long as fishing mortality continues at an
excessive level, the forces affecting "recruitment" make the stocks progressively less
amenable to conventional management and progressively more vulnerable to the
uncontrollable and largely unpredictable physical forces of the environment. In

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1000q

100-
O

UJ
m
10-
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i r i i i i i i i i
25 30 35 40 45 50 55 60 65 70 75
YEAR
FIGURE 6. Numbers of recruits of North Sea haddock determined from catches of 1 -year-old fish by
Scottish research vessels. (From Jones, R. and Hislop, J. R. G., Rapp. Proc. Verg. Res. Cons. Perm.
Int. Expl. Mer., 172:58-71 (1978). With permission.)

short, the "recruitment problem" is like a moving target, and the chances of solving
it seem to be diminishing as heavy fishing pressure persists. There is reason to
believe, however, that the ability to shift along the continuum is not a universal
characteristic of all species. It would appear to be more probable among species at
the midpoint of the continuum and toward the r-end of the spectrum. Strongly
•K-selected species probably have little or no ability to shift toward the r-end of the
continuum. In fact, this inability has been suggested as an explanation for the
extinction of giant reptile species, large pleistocene mammals, or, perhaps, certain
endangered birds (Miller and Botkin, 1974). These examples of the relevance of the
r- and X-selection hypothesis to fishery management anticipate the second and third
components of Margalef s ecosystems dynamics, that is, those of ecosystems devel-
opment and hierarchical structure.

IV THE HYPOTHESIS OF ECOSYSTEMS

DEVELOPMENT
The hypothesis of ecosystems evolution has been developed by a number of
ecologists in varying forms, notably E. P. and H. T. Odum (1969, 1983), Whittaker

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and Woodwell (1972), Margalef (1963, 1975) and Sanders (1968). Table 3 presents
a summary of characteristics of less and more-evolved communities that is of
immediate interest to managers. It indicates that there are different kinds of fish
associations, communities, or systems; that these kinds of communities may be
characterized by the kinds of species found within them; that the yields to be
expected from each are quite different; and that the qualitative response of each to
fishing pressure would be quite different. The time-stability hypothesis of Sanders
(1968) adds a new perspective to the hypothesis of ecosystem development, a
perspective that assists managers in identifying the kinds of communities with which
they are concerned and the kinds of management efforts that are appropriate. It
indicates to us that a young community such as Georges Bank or the Gulf of Maine,
both formed since the most recent glaciation, is probably different in important
qualitative ways from a community in the Gulf of Mexico, that is, different in its
potential yield, its inherent variability, its amenability to or dependence on manage-
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ment, and different in the kinds of species of which they are composed. All these
differences are important to managers.
There is a strong parallel between the kinds of communities and the species
within them (Margalef, 1968). Mature communities are largely composed of equilib-
rium (^-selected) species, whereas the characteristics of immature communities
reflect the large component of opportunistic (r-selected) species. Immature commu-
nities would normally exhibit considerable variability in abundances of species
because each species maximizes reproductive efforts in an environment strongly
influenced by density-independent controlling factors. In contrast, mature commu-
nities in a preexploited state would be relatively predictable, with species fluctuating
only moderately near the carrying capacity.
A few fisheries systems throughout the world are almost fully documented. The
Gulf of Thailand and Georges Bank are two of these. They are undoubtedly
qualitatively different kinds of systems, a fact that becomes apparent when the
general characteristics of each are contrasted.
Two different basic characteristics are immediately apparent in each, that of age
and that of environmental variability. The Gulf of Thailand is literally millions of
years old. Georges Bank is less than 20,000 years old. The temperatures, productiv-
ity, and light characteristics of the Gulf of Thailand are extremely constant or
predictable. The same characteristics of Georges Bank are variable. The concept of
ecosystems development predicts that characteristics for the species in the Gulf of
Thailand would be very different from the Georges Bank. It predicts that an
ecosystem in the Gulf of Thailand with a large component of /C-species, would have
little resilience to accommodate to a major perturbation, such as heavy fishing
pressure. In fact, the fisheries have responded as predicted (Figure 7, Beddington
and May, 1982).
H. T. Odum (1983) makes a distinction between organisms (fish) of lower and
higher embodied energy. (These would be called "trash'' and "market" fish.) The
higher-energy fish have /T-characteristics; the lower-energy, r-characteristics. The
Gulf of Thailand catch data suggest an initial preponderance of /T-species, which
succumbed predictably to fishing effort, and, in addition, a component of more r-
(or less K) selected species, which responded less disastrously to fishing (Figure 8,
Beddington and May, 1982). These data are predictable phenomena from the

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TABLE 3
Ecological succession: trends to be expected in the development
of ecosystems
Ecosystem attributes Developmental stages Mature stages
Community energetics

Gross production/community =1
respiration (P/R ratio)
Gross production/standing crop High Low
biomass (P/B ratio)
Biomass supported/unit energy Low High
flow (B/E ratio)
Net community production High Low
(yield)
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Food chains Linear, predominantly Weblike, predominantly

grazing detritus

Community structure

Total organic matter Small Large

Inorganic nutrients Extrabiotic Intrabiotic
Species diversity — Low High
variety component
Species diversity — Low High
equitability component
Biochemical diversity Low High
Stratification and spatial Poorly organized Well organized
heterogeneity
(pattern diversity)

Life history

Niche specialization Broad Narrow

Size of organism Small Large
Life cycles Short, simple Long, complex

Nutrient cycling

Mineral cycles Open Closed

Nutrient exchange rate, Rapid Slow
between organisms and
environment
Role of detritus in nutrient Unimportant Important
regeneration

<»election pressure

Growth form For rapid growth For feedback control

("r-selection") ("K-selection")
Production Quantity Quality

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TABLE 3 (continued)
Ecological succession: trends to be expected in the development
of ecosystems
Ecosystem attributes Developmental stages Mature stages
Overall homeostasis

Internal symbiosis Undeveloped Developed

Nutrient conservation Poor Good
Stability (resistance to external Poor Good
perturbations)
Entropy High Low
Information Low High
From Odum, E. P., Science, 164:262-270 (1969).
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hypothesis of r- and /^-selection and from Odum's (1983) concept of embodied

energy.
In contrast to the Gulf of Thailand, Georges Bank (Figure 9) indicates notably
different responses by species to fishing pressure (Hennemuth, 1979). There are
variable trends, both up and down, for a number of species. The most prominent
trend, however, was the steady decline of herring, primarily, with apparent compen-
satory increases in several other species. These characteristics of the Georges Bank
response are understandable if it is assumed that the community is a less mature
system than the Gulf of Thailand and is not composed of strongly iC-selected species,
but instead is a mixture of r-selected and moderately JST-selected species, that is, the
species composition occupies a broader segment of the r-K spectrum, with its
statistical mode much closer to the rend than the Gulf of Thailand. The hypotheses
of ecosystems development and of time-stability would predict these community and
species characteristics, given only the brief (<20,000 years) age and the environmen-
tal variability of Georges Bank. The responses of the two systems appear to be in
general conformity with the predictions of the hypotheses. Ursin (1984) noted that
the more specialized a major fishery system is, the more vulnerable it is. The overall
decline of the species of the Gulf of Thailand is, therefore, consistent with Sander's
time-stability hypothesis in that it predicts such specialization.

V. THE CONCEPT OF ECOSYSTEMS HIERARCHY

The third and final component of MargalePs ecosystems dynamics is the mechanisms
that slow the turnover of energy and biomass in systems. These mechanisms are
manifest in biologic communities in general, most having the universal property of
inducing energy and nutrient storage, order, and information transfer, thereby,
building stability and predictability into systems. These mechanisms originated as the
emergent properties of systems to which Dickie referred. They are functions of the
hierarchic constraints on the rates of processes of communities or ecosystems

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 APOLLONIO REVIEWS IN FISHERIES SCIENCE
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SQUID
OTHER GOOD-
SPECIES

PRIACANTHUS
NEMIPTERIDAE
SAURIDAE
LEKDGNATHIDAE
CARANGIDAE

MULLIDAE

SCOLOPSIS

TACHYSURIDAE

RAYS

SCOMBEROMORUS
SHARKS

1963 1966 1968 1970 1972 1974 1976 1978

FIGURE 7. Rsheries landings from the Gulf of Thailand. (From Beddington, J. R. and May, R. M., Sei.
Am., 347:62-69 (1982). With permission.)

(O'Neill et al., 1986). The well-known graph (Figure 10) of Steele (1980a) is a
reflection of this hierarchic structure. Hierarchy has been called a time- and space-
binder because it is an integrating mechanism in both dimensions. As such, it transfers

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 Volume 2 (Issue #2), 1994 ECOSYSTEM CHARACTERISTICS IN FISHERIES MANAGEMENT
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GOOO FISH

TRASH FISH

• 1963 1966 1968 1970 1972 1974 1976 1978

FIGURE 8. Average landings of "good" and "trash" fish from the Gulf of Thailand. (From Beddington,
J. R. and May, R. M., Sei. Am., 247:62-69 (1982). With permission.)

information not only between communities but also into the future (by storing
energy into long-lived, multiple-spawning species) and, thereby, builds predictabil-
ity into systems^ The function of highly migratory, long-lived apex predators is the
most obvious example of space- and time-binders. Migratory species transfer energy
among communities and, just as the movement of air masses stabilizes climates, in
the same manner migrants tend to stabilize marine communities. Whale migrations
move species from species-rich to -poor communities, and they move energy and
nutrients across the rich-to-poor gradients of these systems. Although these are
obvious and dramatic examples of hierarchic function in the sea, there can be little
doubt that the hierarchic principal also works at less obvious, more subtle levels.
Steele (1980b) described parallels between a hierarchy of physical océano-
graphie features and a hierarchy of species aggregations or "patchiness" through
trophic levels from phytoplankton to Zooplankton to herring (Figure 11). The
progressive hierarchy of those aggregations was verified by statistical analyses for
a number of species by Hennemuth and Shepherd (1983). Steele concluded that the
progressive degree of aggregations reflected both an increasing social structure and
an increasing independence from control by physical factors; the hierarchy pro-
gressed from control by physical forces on the lower strata of species to significant
uses of physical forces by higher species. Further, Steele's (1980b) diagram of
aggregations conveys the essentials of biologic controls within the hierarchic
structure, that is, the higher strata are closely tied to the resource. Their prey, in turn,

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 APOLLONIO REVIEWS IN FISHERIES SCIENCE

10« -

/V r^^\ '-TOTAL
A

TV /» / • — ^

(v A/Z\A \WMACKEREL
10'

/\^ // yY/ \ \
\ \4, /^^
/
/^>y
\ /
/X-^
vl
Y<J-- OTHER FWFBH
\ SUVER HAKE
^—v
Downloaded by [New York University] at 19:08 05 June 2015

' \ - ^
4/ // / . AV\ r/\ V VSQOO
>r / y ^/ i P r C R ^ ^ - ^ ^ T ^ OTHER X^ \ W ^ * C — ^«s\ FLDUNOeR
W/»» \ U^-POLUDCK
>
7/r V ^ \V~-REDF1SH
4
s/\ £ N \V-HADOOCK
^•\/f \ V-REDHAKE

V \ / V /»-YTflOUMJER

10' .A 1 V /

GEORGES BANK
TOTAL LANDINGS
m* i i i i t 1 1 1 1 1 i i i i i i i i i i i i i
19S5 57 59 61 63 65 67 69 71 73 75 77
YEAR

FIGURE 9. Total fisheries landings from Georges Bank. (From Hennemuth, R. C , Contemporary
Quantitative Ecology and Related Ecometrics, In. Coop. Publishing, 1979. With permission.)

are substantially impacted by prédation. Lower in the hierarchy, prédation (biologi-

cal control) is progressively less significant, and physical control is increasingly
dominant. Steele's observations on aggregations makes particularly clear one method
of ecosystems control through hierarchic structure. Other kinds of hierarchies would
be equally important, some of which may not yet be defined.

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 Volume 2 (Issue #2), 1994 ECOSYSTEM CHARACTERISTICS IN FISHERIES MANAGEMENT

îooo r

100

K)

1.0-

1.0 10 100 1000

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KILOMETERS

FIGURE 10. Spatial and temporal scales of phytoplankton [P], Zooplankton [Z], and fish [F]. (Modified
from Steele, J. H., Natl. Acad. of Sei., 1980a. With permission.)

FRACTION OF TOTAL VOLUME

FIGURE 11. Schematic, and hypothetical, instantaneous distributions depicting the relation between
concentration of three trophic levels and the fraction of the upper layers of the ocean they may occupy.
(From Steele, J. H., Oceanus, 23:3-8 (1980b). With permission.)

An extended review of this hypothesis (O'Neil et al., 1986) makes the essential
point that the fundamental parameters of ecosystem hierarchies are constraints of
process rates, that is, rates of energy flow, nutrient recycling, metabolism, fecundity,
growth, prédation, maturity, mortality, and generation. This principle is recognizable
to managers when the relation of apex predators to their prey is considered. The
organization of ecosystems is a consequence of hierarchic differences in process
rates, the higher levels exerting a regulatory constraint over the rates in lower levels.
These differences in rates are in fact the essence of the significance of distinction
among r- and /T-selected species. These species-rate differences exert control over
systems properties and functions through a hierarchic structure.

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 APOLLONIO REVIEWS IN FISHERIES SCIENCE

This concept of linking species characteristics to the mechanisms of ecosystem

structure by rate processes is the answer, in principle, to how the system works. It
is this concept that is essential for managers to understand.
Managers should be aware of this hierarchic principle because the disruption of
hierarchy, the mechanism that regulates process rates, results in less predictability
in a community, and in less manageability of the system.

VI. MANAGEMENT IMPLICATIONS OF ECOSYSTEMS

CONCEPTS
Ecosystems and the species of which systems are comprised evolve with tenden-
cies predictable from the three basic hypotheses discussed in the previous
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sections. Fishing efforts, in general, reverse that natural evolution of species,

systems, and organizing mechanisms. This reversal is partly by design, but many
of the inevitable consequences are not, and they are not desirable from a
managers' perspective.
Conventional fishing strategy is to maximize the productivity of the biomass
of individual species and to induce a fishing effort that maximizes the yield per
recruit. This is contrary to tendencies of natural evolution toward increased
efficiency of resource use and, thus, a high ratio of biomass to production.
Associated necessarily with an increase of the ratio of production to biomass as
a result of conventional fishing strategy are several other realities, most are familiar
to fisheries managers. These are decreased year classes and decreased spawning
periods; decreased age at maturity and increased growth rates (both compensatory
responses to fishing pressure); and, less obvious, reduction of innate biologic
feedback mechanisms that are usually associated with older-year classes of a species.
All of these are symptoms of a species shift toward greater r-selectivity and lesser
community organization (Margalef, 1968). The conventional fishing strategy assumes
that fishing effort can be controlled at the point of maximized yield-per-recruit.
Unfortunately, this rarely happens. Experience shows that fishing effort routinely far
exceeds the desired point, and, as a result, yield per recruit declines. Other life
history parameters also are inevitably affected, resulting in the species necessarily
acquiring r-selected characteristics, that is, enhanced instability and unpredictability,
greater vulnerability to Oargely unpredictable) physical environmental perturbations,
and less value in the market Beddington and May (1977) concluded that, as fishing
mortality (iO increases toward the fishing mortality at maximum sustainable yield
ÖVisyX variability in population biomass increases — a qualitative change that is
predictable from basic theory as a consequence of fishing strategy. All of this is
inevitable and is confirmed by typical management experience. More importantly,
it is predictable from basic ecosystems theory, concepts that may not be familiar to
managers.
Further, the species associations or the fish communities as a whole are similarly
affected, a fact that may also be predicted. As the communities reflect the charac-
teristics of the species comprising them, and as species are forced by unrestrained
fishing effect toward r-selection, in the same manner the community is forced toward
lesser maturity. The predictions of the ecosystems hypotheses are supported by New

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 Volume 2 (Issue #2), 1994 ECOSYSTEM CHARACTERISTICS IN FISHERIES MANAGEMENT

England experience, that is, as the high-value species have been fished down,
increasing attention has been focused on species of lower values, such as squid. As
high-value species in the North Sea were depleted, there appeared an abundance
of pout, sand eels, and anchovies — industrial species. As Pacific ocean perch were
depleted, there was an abundance of pollock. Regier (1973) documented a shift of
the species composition of the fish community of Lake Erie under fishing and
écologie stress: (1) replacement of large benthic species of high value by small
pelagic species of lower market value, (2) increased variability in year-to-year
landings. All these effects are consistent with predictions.
It should be noted that species may be destabilized in the sense of increasing
their relative r-selectivity by reduction of the numbers of year classes in the
population. Communities may be forced toward lesser maturities by a number of
mechanisms: (1) by forcing their component species toward r-selection, (2) by
virtually eliminating species as significant components of the system, (3) by changing
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the relative abundance (and the balance of r- and /T-selection) of remaining species,
and (4) by focusing fishing effort at lower levels of the trophic structure, as in the
krill fishery. There is a cumulative impact of all these forces on the community.
Mechanized fishing does all these things simultaneously; in short, it has the inherent
effect of making stocks progressively unmanageable.
In simple but useful terms, how the system works is summarized in Figure 12.
The contrary tendencies of natural evolution on the one hand and of fishing on the
other on the system dynamics are apparent, as is also the disparity in time of
response. Fishery managers are not likely to witness a natural evolution of a system.
Their interest in that context is simply an insight into the kind of community under
their jurisdiction, based upon its longevity. But most managers have experienced or
may anticipate seeing the reversal ofthat evolution, which may occur, perhaps, with
a rapidity that challenges their management capabilities.
This concept of a few basic hypotheses underlying the nature of fisheries
ecosystems has several useful features:

1. It conforms in principle to the practical experience of managers and of

fishermen.
2. It is consistent with the history of many fisheries.
3. It appears to explain why those trends have developed as they have.
4. It predicts what will happen to various kinds of stocks under continuous
fishing pressure.
5. It urges that managers pay particular attention to the kinds of fish species and
fish stocks under their jurisdiction.
6. It reconciles divergent views; for example, the assumption of an inherent
equilibrium in fisheries as opposed to a constant change in fisheries — both
views can be accommodated at appropriate positions on the continuum of
species and community characteristics.
7. It offers an explanation of why fisheries models work in some instances and
not in other.

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 APOLLONIO REVIEWS IN FISHERIES SCIENCE

EVOLUTION OF COMMUNITY DYNAMICS

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A. Direction of natural evolution

( 1 0 5 - 1 0 7 Years)

B. Direction of fishing perturbations

( 1 0 ° - 1 0 1 Years)

FIGURE 12. Summary of the evolution of community dynamics under (A) natural evolution and (B)
fishing perturbation.

8. It conforms to the widely accepted views of the way species, communities,

and fisheries in general evolve throughout the world.

It would be most useful if the concept predicted the consequences of various

kinds of management; however, the outlook about the future is never as clear as the
view of the present or the past. However, these hypotheses are useful for manage-
ment guidance.

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 Volume 2 (Issue #2), 1994 ECOSYSTEM CHARACTERISTICS IN FISHERIES MANAGEMENT

VII. MANAGEMENT BASED ON ECOSYSTEMS

PRINCIPLES
First, the most obvious lesson is that if fishing vessels are to replace apex predators,
in a sense they have, then they must have characteristics comparable to apex
predators if the systems are to be manageable, that is, the vessels must emulate the
essential characteristics of ÜT-selected species. Appropriate feedback mechanisms
between vessels and prey species would be most important. These are presently
almost totally absent, particularly in the case of pelagic-species purse seiners.
Subsidies to vessels is a way that reverses normal feedback mechanisms. A multispecies
vessel switching rapidly from one species to another is another way — both working
contrary to the normal, narrow niche specificity of apex predators. Tax write-offs and
investments from outside the industry, such as joint ventures, also, are destabilizing
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features of the industry.

A further example is the lack of regulating linkages between resource conditions
and the processing and marketing of product. Herring processors in Maine or scallop
processors in New Bedford are mildly inconvenienced by shortages of local fish;
however, they continue to prosper by importation of raw material from northwestern
Europe or from around the world. Their success is not linked to the condition of
the local stocks; thus, they feel no urgency for their effective management.
Similarly, market forces provide no effective feedback mechanisms from the
consumer through the domestic vessels to the local stocks. On the contrary, imports
induce positive, nonregulatory feedbacks. Consumers show little or no preference
for the source of their fish. Thus, imports are able to sustain a supply that satisfies
growing demand and supports rising prices. The local vessel perseveres on depleted
stocks because of rising prices and, thereby, is less interested in their effective
management as long as there remains any resource at all to harvest.
Industry practices and economic forces work against the innate stabilizing
mechanisms of naturally evolved, preexploited systems. Managers, therefore, must
recognize that their regulations must balance — indeed, must overcome — a variety
of powerful, destabilizing forces.
Second, the hypotheses indicate that simple reduction of fishing effort may not
be sufficient for rebuilding the stocks to an abundance and composition similar to
those of preexploitation days. Under reduced fishing pressure, the opportunistic
species might simply outcompete the slower-growing, more valuable, more JT-selected
species and, thus, dominate the biomass. However, this event means continuous
large fluctuations of species abundances and little manageability inherent in the
system.
Third, hierarchy theory (O'Neill, et al., 1986) suggests that restoration of
traditional stock composition would be attained only by well-defined management
efforts; these might possibly include maintaining a high fishing effort on certain
species in imitation of predator species, until such time as predators have rebuilt not
only stock abundances but also age structures that induce stabilizing mechanisms
into the system. The difficulty with this possibility is that there may be more
r-selected species to be held in check than may be practicable. More subtle, more
carefully contrived measures are probably essential.

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Fourth, the ecosystem concept indicates that the biological attributes of the
species, which, to a degree, we have neglected, must be taken into account and that
restoration of the structure and manageability of communities will be based on
biologic knowledge of the structuring components of the system.
Fifth, the concept indicates that all components cannot be maximized simulta-
neously.
The challenge is to convert a set of ideas that appears relevant to fisheries to
hypotheses that are useful to their managers. That challenge is particularly important
for the question of how it would impact on effective management. Although the
ideas are persuasive in forecasting the effects of continuous fishing, the greater
challenge is to forecast the outcomes of management.
The essential overriding virtue of these ideas is that they offer the possibility of
a cohesive understanding of how systems may be identified and how systems
behave. They offer a unifying and simplifying view of fisheries; science progresses
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by the simplification and unification of ideas and observations through the formu-
lation of fundamental hypotheses.
If these or similar ideas do not form such hypotheses, one must ask why
fisheries are fundamentally different from other ecosystems. Why do rules that apply
elsewhere not apply in the sea? And if they, in fact, do not, then an alternative set
of hypotheses that better explain our observations should be formulated. It is only
within some such hypotheses that ecosystem management can be effective and
productive.

ACKNOWLEDGMENT
I thank Lloyd M. Dickie for suggestions for several notable improvements to the
article.

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