or actual animal conflicts in social foraging
(among other topics), and a verbal consid-
eration of the applicability of game theory
to important aspects of human behaviour
such as social norms.
The common framework for the selec-
tion of papers is classic evolutionary game
theory – perhaps one of the most influential
paradigms of the century – initiated by
Von Neumann and Morgenstern
2
, Nash
3
and Maynard Smith
4
. This equilibrium-
oriented approach proves to be as useful as
expected from past and recent develop-
ments in the discipline. Realistic, ‘field-
motivated’ situations of the many different
types of animal conflict covered in the book
fit nicely into elegant, mathematically sim-
ple models. Most of the authors compare
model predictions with sound empirical
evidence – although the amount of evi-
dence and taxonomic diversity discussed
in each chapter are quite different. One has
the impression that empirical work keeps
pace with game-theoretical modelling in the
study of animal behaviour, which is an envi-
able state of affairs compared with some
other disciplines of theoretical biology.
The general applicability of classic evo-
lutionary game theory to problems far from
those of animal conflicts is a recurrent
claim of publications on the subject. The
usual list of disciplines taking advantage
of such secondary applications includes
economics, social science and even the
humanities. Interestingly, it is rarely stated
that evolutionary game theory has much to
say to other fields of population biology.
For example, within-community multi-
species coevolution of plants calls for a
game-theoretic approach, with growth
forms and/or dispersal techniques consid-
ered as strategies, and possible stable
equilibria representing evolutionarily sta-
ble coalitions. There are good reasons to
expect that game theory will help us
explain plant ‘behaviour’ and, on that basis,
certain aspects of community diversity.
This would be yet another link in the
promising relationship between game theory
and evolutionary ecology.
Tamás Czárán
Ecological Modelling Research Group,
Hungarian Academy of Science and Eötvös
University, Budapest, Hungary
(czaran@ludens.elte.hu)
References
1 Durrett, R. and Levin, S.A. (1994) Theor. Popul.
Biol. 46, 363–394
2 Von Neumann, J. and Morgenstern, O. (1944)
Theory of Games and Economic Behaviour,
Princeton University Press
3 Nash, J.F. (1951) Ann. Math. 54, 286–295
4 Maynard Smith, J. (1982) Evolution and the
Theory of Games, Cambridge University Press
Rivers of life
Restoring Life in Running
Waters: Better Biological
Monitoring
by J.R. Karr and E.W. Chu
Island Press, 1998.
$29.95 pbk (xiv + 206 pages)
ISBN 1 55963 674 2
T
he concept of ecological services has
been warmly embraced as a pragmatic
justification for the conservation of biologi-
cal diversity
1,2
. Experiments reveal that
more diverse terrestrial communities are
better at fixing carbon dioxide (CO
2
) or
show increased resilience in the face of
drought or other perturbations
3,4
. Unfortu-
nately, the generality of these findings is
unclear because the experimental commu-
nities involved are, of necessity, small or
artificially contrived, and the challenges of
extrapolating the results, to rain forests for
example, are considerable. In freshwater
systems, by contrast, the logic for conser-
vation is incontrovertible. Clean water is
fundamental to life. Healthy river systems
deliver not just a water supply, but also
fish, power generation, transport, recre-
ation and cultural identity. The demand for
these services will intensify as populations
grow and countries develop, and it is
widely predicted that the battles of the 21st
century will be waged over water resources
rather than oil or territory.
Biologists are faced with the challenge
of monitoring the wellbeing of freshwater
habitats. Happily, the task is relatively
straightforward because the status of the
living organisms found there is the best
indicator of condition. Even more propi-
tiously, the preservation or restoration of
natural systems, and the associated biologi-
cal diversity, emerges as the most effective
way of guarding freshwater resources. With
this impetus, it is not surprising that con-
siderable effort has been devoted to creat-
ing measures that encapsulate biological
quality in a single figure. Karr and Chu
review this approach and make a strong
case for the adoption of multimetric biologi-
cal indices in river assessment and manage-
ment. Although their book is focused on the
United States, much of the discussion is of
wider relevance.
Multimetric measures are used to gen-
erate an index of biological integrity (IBI).
This is a composite of around eight to 12
metrics, or variables, each chosen to reflect
some aspect of the system’s biological
health. The metrics span a range of attrib-
utes, including the richness of specified
taxa, trophic structure, such as the relative
abundance of predators, and rates of dis-
ease or malformation in target species. They
need not be independent. Indeed, a typical
IBI, such as one used in the midwestern
USA, includes the total number of fish species
as metrics, as well as the richness of sep-
arate taxonomic groupings, such as darters
and sunfish. Each metric is ranked from one
to five, with five being the value expected
for a pristine habitat. The metrics are
summed to produce the final index, which
is then used to compare and assess sites.
Ecologists are often suspicious of meas-
ures that pool disparate variables. Experi-
ence with diversity statistics, such as the
Shannon index, has for instance revealed
how an assumed virtue – in this case, the
integration of richness and evenness
scores – can thwart interpretation. Nonethe-
less, the authors discuss 37 premises (e.g.
‘statistical decision rules are no substitute
for biological judgement’) on which the
case for multimetric measures is based,
and criticize seven ‘myths’ (e.g. ‘the sensi-
tivity of multimetric measures is unknown’)
that have been advanced against them.
Karr and Chu also note that multimetric
indices are statistically versatile and can be
analysed with familiar techniques, such as
ANOVA or regression, and that the preci-
sion of sampling protocols can be esti-
mated in conventional ways. Moreover, IBIs
are not just confined to rivers. The book
lists a variety of environments, including
lakes and coastal and marine systems, in
which they have been successfully applied.
Karr and Chu argue that multimetric
measures are easily comprehensible and
draw an analogy with economic indicators
such as the Dow Jones industrial average. But
whereas the Dow Jones generates a single
measure of the economic status of one coun-
try, IBIs are faced with a broader task. Pris-
tine freshwater habitats can vary markedly
from place to place and estimates of species
richness (a central component of many
IBIs) are not an infallible guide to ecological
quality. Headwater streams, for example, typi-
cally support fewer species than large rivers.
The authors address this concern, maintain
that multimetric measures can be wide-
ranging and cite, as evidence, a fish IBI that
successfully classified sites in six US
regions ranging from Chicago to Arkansas.
However, they also recognize that the proper
classification of sites, so that like is com-
pared with like, is the key to a successful IBI.
There is no doubt that IBIs have been a
valuable tool in the monitoring and conser-
vation of American rivers and Karr and Chu
provide a compelling case for their univer-
sal adoption. Given the importance of
benchmark data in the selection of metrics
and appropriate deployment of IBIs, it
remains to be seen if they can fulfil their
promise in diverse, and increasingly threat-
ened, tropical rivers for which we have
depressingly little biological information.
BOOK REVIEWS
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