PERSPECTIVES

ously more tractable than the logarithmic

What does ‘biodiversity’ mean - series.
Later, an interesting model was pro-
scientific problem or convenient myth? posed by MacArthur-h,who assumed that
boundaries between niches in resource
niche hypervolume are set at random,
whereas the relative abundances of spe-
Alexej Ghilarov
cies are proportional to these sections of
hypervolume. This model became widely
Biodiversity has become an everyday word, not just for ecologists but for politicians,
known as ‘broken-stick, or MacArthur’s
the media and the general public too. But does It have a genuine scientific meaning
model. The distribution of abundance pre-
that distinguishes it from earlier studies of diversity, or has It assumed a
scribed by MacArthur’s model is much
mythological status?
‘flatter’ (i.e. the contrast between given
Alexej Ghilarov is at the Dept of Vertebrate Zoology and General Ecology, species and the next in the sequence is
Biological Faculty, M.V. Lomonosov Moscow University, less) than in the case of a logarithmic
Moscow 119899, Russia. series.
In the late 1960s and early 197Os, it
became clear that there is no universal
he term ‘biodiversity’ appeared in eco- much more appropriatei. At that time, type of distribution of relative abundance
T logical literature in the middle of the
1980s and in less than 10 years it became
Fisher worked at Rothamsted Experimen-
tal Station (UK), where another ecologist,
that corresponds to all real communities,
though such distributions change in
so popular that for an external observer C.B.Williams, had the results of four years the course of succession according to a
the whole situation looks like a successful of continuous catching of Lepidoptera in a particular pattern. The dominance of a few
breakthrough in some new field of science. light trap. This huge sample (which in- of the most-abundant species is more pro-
It seems, however, that the real reason for cluded more than 16 000 moths identified nounced at the early stages of succession,
this burst of interest in biodiversity (or to about 240 species) was truly random while later, the species of intermediate
rather this rapid increase in the usage because the trap did not cease to collect abundance become more significants.How-
of the term ‘biodiversity’) lies outside the any extra individuals of any given abundant ever, it seems that a deep understanding
sphere of science. To many professional species. In their joint paper, published in of underlying mechanisms that result in a
ecologists, perhaps it sounds offensive, 1943 in The Journal of Animal Ecology, given pattern of species abundance has
yet I shall dare to claim that the term ‘bio- Fisher, Corbet and Williams* proposed to not been achieved.
diversity’ became so popular, primarily be- approximate the frequency distribution of Another line of species diversity stud-
cause its usage facilitates greater access the species represented by 1,2,3,4,.. . (and ies in the 1960s and 1970s was connected
to research funding. So, we might consider so on) individuals by a logarithmic series: with the use of special indices proposed
‘biodiversity’ as a useful fad that helps ox, ti2/2, 0x3/3, (wx4/4,..., where the con- to measure diversity without reference to
many ecologists to survive doing science. stant, cx,has been found to be a measure some hypothetical distribution of relative
However, by characterizing something of species diversity. Species diversity is abundance. A great variety of indices was
as a ‘fad’we simply dodge the question low when the number of species is grow- proposed that assess the number of spe
of why ‘biodiversity’ has caught on and ing slowly with the increase in number of ties and the proportions in abundance of
other terms have not. It is clear that more individuals, and high when the number different species. Among others, there was
thorough analysis of the whole situation is of species is growing quickly. the very popular index that is based on
needed. First, we must try to understand Thus, the problem of species diversity Shannon’s formula derived from infor-
to what extent the rapidly growing usage of measurement was not only clearly formu- mation theory:
the very word ‘biodiversity’ reflects a real lated more than 50 years ago, but, to some
H = -C/I, log p;
increase in knowledge of all the phenom- extent, it was resolved by proposing a par-
ena called by this name. Second, we must ticular index. It was also important that where pi is the proportion of the total
discuss the reasonable borders of broad- Fisher et al. attempted to find some gen- number of individuals that belongs to the
ening the context of biodiversity. In par- eral ‘rule’or ‘law’,according to which the ith species.
ticular, we may ask if it is necessary to numerical abundances of different species It was probably MargalefG who first
justify the studies of biodiversity by refer- were related to each other. In emphasiz- used this index (though expressed in a dif-
ring to the possible interrelations between ing the importance of Fisher etal.3 paper, ferent form) and who proposed to evalu-
community diversity and ecosystem ‘func- we must point out that its publication ate the level of community organization in
tioning’. Third, we may ask ourselves what was not an isolated event. In 1948,Preston3 terms of information theory. The writings
the growing popularity of ‘biodiversity’ showed that in many communities, the of Margalef stimulated many ecologists of
means for ecology. number of species with given abundance the 1960s to measure quantitatively the
could be approximated by the lognormal species diversity of different communities
Past achievements distribution. If species are classified in and/or of the same community in different
Diversity is not a new theme for ecolo- accordance with their abundance in loga- stages of its development. At that time,
gists. In the early 1940s A.S. Corbet, ana- rithmically increasing classes - so-called there was a widespread belief (though hid-
lysing a large collection of butterflies from ‘octaves’ (i.e. the first octave contains l-2 den rather than clearly formulated) that
Malaya, remarked on the decrease in num- individuals, the second, 2-4 individuals, with a single numerical value, we can as-
ber of new species with an increasing num- the third, 4-8, the fourth, 8-16, and so on) sess some very significant feature of com-
ber of individuals. He supposed that the -then the number of species per ‘octave’ munity structure. It was also important
resulting distribution could be described shows a truncated normal distribution. If that such studies were done just at the
by a hyperbola, but R.A. Fisher, to whom a sample contains a high number of spe- . community level. Therefore, many ecolo-
Corbet sent his results, suggested that a ties and individuals, we can usually obtain gists believed that in measuring species
negative binomial distribution would be a lognormal distribution, and it is obvi- diversity they were using a methodology

304 0 1996, Elsevier Science Ltd Pll: SO169-5347(96)20016-3 TREE vol. II, no. 7 July 1996

that was in accord with the very old but
ever-popular idea of the community as
something more than a mere sum of separ-
ate populations. Thus, it was a pleasantly
familiar idea.
Ecologists of the 1960s and 1970s did
not, however, confine themselves to meas-
uring species diversity and the simple ac-
cumulation of such descriptive data. They
went further, in trying to understand the
relationship of diversity with other fea-
tures of community, and to consider the
whole problem in a broader context (see,
for instance, the reviews of Williams’,
MacArthur7 and Whittakers). Special at-
tention was drawn to the dependence of
species diversity on the structural com-
plexity of the environments, to the role of
predation9 or periodical disturbance’0 in
determining a given level of diversity, and
to the relationship between the species di-
versity and standing crop of communityli.
Thus, it seems that although biodiver-
sity might be a new word, it is not a new
topic for ecology. Moreover, considerable
progress in the study of species diversity
had already been achieved 25-30 years
ago. In the past decade, this line of studies
attracted less interest, but it was not cut off
completely. I suppose that the reasons for
such changes are determined by the inner
logic of ecology’s development and its turn
toward a more reductionistic approach,
rather than by the influence of some ex-
ternal forces.
The ecosystem and the aeroplane
According to Harper and Hawks-
worth”, the expression ‘biological diver-
sity’ was used in 1980 by Lovejoy and
Norse and McManusi4. If for Lovejoy, it
meant primarily the number of species,
the latter authors used it to designate both
genetic and ecological diversity. By 1981,
the ‘USStrategy Conference on Biological
Diversity’ had taken place, while in 1985,
W. Rosen used the contracted form ‘bio-
diversity’ for planning the conference
‘National Forum on BioDiversity’that took
place in Washington, DC, in September
1986. As Harper and Hawksworth re-
marked’*, ‘The proceedings of that forum,
edited by Wilson...under the title Bio-
diversity, launched the word into general
use.’ In 1992, the official representatives
of more than 50 nations signed the ‘Con-
vention on Biological Diversity’, which can
be considered as official recognition of the
importance of biodiversity.
It is worth noting that from the very
beginning, the use of ‘biodiversity’ was
connected with politics and environmen-
tal technology rather than with the science
itself. However, scientists tried to exploit
the popularity of this term for their own
sake-to get money to enable the continu-
ation of their previous work (mainly in the
TREE vol. II, no. 7July 1996
sphere of systematics and ecology); work
that seemed related to the problems of bio
diversity. Sometimes, even serious scien-
tists were obliged to contribute to the
myth of biodiversity by showing its practi-
cal meaning for a broad public. That might
be done in two ways: first, by emphasizing
the direct value of biodiversity considered
as the variety of potential biological re-
sources that can be used in pharmacy, agri-
culture or for other purposes; and second,
by addressing the possible role of bio-
diversity as a necessary condition for eco-
system functioning. Developing the latter
approach, Paul and Anne Ehrlichis pro-
posed the analogy of the ecosystem with
the aeroplane - the diversity of living forms
representing rivets; each plays a small but
significant role in the working of the
whole. This comparison of the aeroplane
with the ecosystem became rather popu-
lar despite (or perhaps because of) its
extreme superficiality.
An aeroplane certainly needs some
quantity of rivets to fly, and an increase
in the number of rivets can diminish the
probability of a crash, but very soon this
dependence levels off. Therefore the re-
lation between the number of rivets and
plane safety has a distinct threshold effect.
The plane that can move on the ground but
cannot fly is not a plane at all. However, if
somewhere on the earth some organisms
are living, we can be sure that they are the
components of some ecosystem (actually,
it is a simple consequence of the eco-
system’s definition). While it is obvious
that plane ‘functioning’ is flying, the defi-
nition of ecosystem ‘functioning’is rather
vague. Usually it is tacitly admitted that
ecosystem functioning means the pro-
cesses of production and/or destruction of
organic matter, as well as cycling of vari-
ous elements.
In contrast to the plane, which is a dis-
crete object with obviously designated
borders, the ‘volume’of the ecosystem is
not definite, varying in its scale depending
on the process that is used to define it.
Thus, using the completeness of phospho
rus cycling as the criterion of a freshwater
ecosystem, we can consider the epilimnion
of a mesotrophic temperate lake as a good
ecosystem if the period of our obser-
vations embraces several weeks during
summer stratification. However, ifwe extend
the period of study to several months, so
that the time of autumn mixing is included,
only the whole water column can be
called the ecosystem. If the investigation
covers the entire year, the income of melt
waters must be taken into consideration,
and perhaps not only the waterbody itself
but also the entire watershed must be
considered as a part of the ecosystem.
Another problem concerns ecosystem
development and the reliability of com-
PERSPECTIVES
parison of different ecosystems. Even a
very primitive ecosystem (e.g. a newly ex-
cavated pond, or a patch of cleared for-
est) is already an ecosystem that can be
characterized by some structure and func-
tioning. Later, rather complex ecosystems
of high species diversity can develop in
these sites. However, the question arises
whether or not we can show the meaning
of species diversity for ecosystem func-
tioning by comparing different intermedi-
ate stages of ecosystem development.
For a long time, a regression model
that described the dependence of phyto-
plankton biomass in lakes on phosphorus
loading was considered to be an excellent
example of a predictive (and understand-
able!) cause-effect model. However, as
Harris16 recently remarked, this relation-
ship appears true only because of the sig-
nificant differences in organization and
functioning between oligotrophic and eu-
trophic ecosystems (in the former, strong
grazing pressure upon phytoplankton rep
resented by small organisms contributes
considerably to nutrient recycling, while
in the latter, grazing is negligible but the
external input of nutrients is very impor-
tant). Therefore, both waterbodies that are
embraced by the one regression model
cannot actually be considered to differ only
by the degree of quantitative expression
of the same property, and the results of the
model do not illustrate the mechanism
that seems to underlie the relationship be-
tween nutrient loading and phytoplankton
abundance.
In a similar way, the comparison of
functional properties of ecosystems that
are characterized by different levels of
their species diversity does not necess-
arily demonstrate the importance of di-
versity for ecosystem functioning. Simple
solutions to these rather difficult prob-
lems are not necessarily correct. Thus, the
recent attempt by Naeem et a/.17 to ap-
proach the relationship between diversity
and productivity by direct measurement
of aboveground biomass of annual plants
seeded in different combinations looks
brave but optimistic, despite ‘positive’ re-
sults of these experiments (see also the
commentary of Marcel et ~1.18).
Another aspect of the ‘rivet hypothesis’
concerns its use in the practice of conser-
vation. Though it is easy to declare the im-
portance of all species for successful eco-
system functioning, it is hardly possible to
show this importance practically. In our
estimates of ‘functioning’, we usually refer
to such ecosystem properties that resulted
from additive individual activity of many
specimens (e.g. their photosynthesis, res-
piration, excretion or growth). Therefore,
rare species, even if they are represented
by organisms of large size, contribute neg-
ligibly to ecosystem functioning, and their
305
PERSPECTIVES

removal usually (but not always!) does not
significantly influence the rest of the biotic
community. Thus, the bear (Ursus arctos)
was a common species in forests just near
Moscow up to the 17th century. Now, un-
fortunately, there are no more bears in the
Moscow region, but the forests continue
to grow, consuming carbon dioxide and
releasing oxygen in the same manner as
they did centuries ago.
Inevitable mythology
In contrast to mythology and technol-
ogy, which are the universal products of
development of human societies, emerg-
ing in some periods in all known civiliz-
ations, science is unique. Born 25 centuries
ago in the Eastern Mediterranean, science
developed in accordance with its own
methodology, which is rather strict, and
obligatory for everybodylg. Unfortunately,
even in developed countries, scientific
thinking and the scientific view of reality
are characteristic only of a small minor-
ity of people, while the mythological ap-
proach, being more ancient and more uni-
versal, is still alive and widespread. It plays
some hidden role even in sciencezo, not to
mention the sphere of politics and the
everyday life of many people. Technology,
being the form of human activity inter-
ested only in final results but not in the
ways of their achievement (though means
are important in terms of cost), coexists
with mythology but does not necessarily
contribute to the development of science.
Foundation for Basic Research, and
partly by a grant from the state program
‘Biologicheskoe raznoobrazie’.
References
1 Williams, C.B. (1964) Patterns in the Balance of
Nature, Academic Press
2 Fisher, R.A.,Corbet, A.S. and Williams, C.B.
(1943) The relation between the number of
species and the number of individuals in a
random sample of an animal population,
J. Anim. Ecol. l&42-58
3 Preston, F.W. (1948) The commonness and
rarity of species, Ecology 29,254-283
4 MacArthur, R.H. (1957) On the relative
abundance of bird species, Proc.Nat1Acad.
Sci. USA 45,293-295
5 Whittaker, R.H. (1972) Evolution and
measurement of species diversity, Taxon 2 1,
213-251
6 Margalef, R. (1957) La teorla de la lnformaci6n
en ecologia, Memorias de la Real Academia de
Cienciasy Artes (Barcelona), 3rd ser. 32,373-449
7 MacArthur, R.H. (1965) Patterns of species
diversity, Biol. Rev. 40,510-533
8 MacArthur, R. and MacArthur, J. (1961) On
bird species diversity, Ecology 42,594-598
9 Addicott, J.F. (1974) Predation and prey
community structure: an experimental study
of mosquito larvae on the protozoan
communities of pitcher plants, Ecology 55,
475-492
10 Sousa, W.P. (1979) Disturbance in marine
intertidal boulder fields: the nonequilibrium
maintenance of species diversity, Ecology60,
1225-1239
trends b
11 Ghilarov, A.M. and Timonin, A.G. (1972)
Relations between biomass and species
diversity in marine and freshwater
zooplankton communities, Oikos 23,
190-196
12 Harper, J.L. and Hawksworth, D.L. (1994)
Biodiversity: measurement and
estimation, Philos. Trans. R. Sot. London Ser. B
345,5-12
13 Lovejoy, T.E. (1980) The Global 2000
Report to the President (Vol. 2) (The Technical
Report) (Barney, GO., ed.), pp. 327-332,
Penguin
14 Norse, E.A. and McManus, R.E. (1980)
Environmental Qualily 1980: The Eleventh
Annual Report of the Council on Environmental
Quality,31-80, Council on Environmental
Quality
15 Baskin, Y. (1994) Ecologists dare to ask How
much does diversity matter? Science 264,
202-203
16 Harris, G.P. (1994) Pattern, process and
prediction in aquatic ecology. A lbnnological
view of some general ecological problems,
FreshLu.Biol. 32,143-160
17 Naeem, S. et al. (1994) Declining biodiversity
can alter the performance of ecosystems,
Nature 368,734-737
18 Marcel, A., Br&hignac, F. and Thibault, P.
(1994) Blodlversity in model ecosystems,
Nature 371,565
19 Levin, A.E. (1977) Myth, technology, science,
Priroda 3,88-101 [in Russian]
20 Ghilarov, A.M. (1991) Ecology, mythology and
the organismic way of thinking in lhnnology,
Trends Ecol. Evol. 7,22-25

ECOLOGY&
The expense of military pursuits world-
wide exceeds many times over the funds
that are diverted to science and education
worldwide. Science has to disguise itself as
technology, and/or even as mythology, in
EVOLUTIOIV
order to procure the money for subsistence.

Coming soon in TREE

The very term ‘biodiversity’ is an element
of this disguise. This specially invented word
sounds like a proper noun, but it is well
known that proper nouns are especially Quantitative genetics and the assessment of
characteristic of mythological thinking. variation in endangered species,
And last but not least: in challenging
scientists to do science (and not to do A. Storfer
mythology) I do not wish to imply that
The conservation value of fragments of
biodiversity does not require our efforts to
be conserved. Certainly,we must do our best lowland tropical rain forest,
to defend all forms of life, including the gen- Turner and R. T Corlett
etic diversity of populations, the diversity of
populations and species, as well as the di- Sources and sinks in population biology,
versity of communities and landscapes. I P.C. L&s
suppose that justifications from the spheres
of ethics and aesthetics must be used much Ecological immunology: costly parasite deft!nces
more broadly. Where science is concerned, and tradmffs in evolutionary ecology,
reasoning must develop according to inner
logic, and science must solve primarily B.C. Sheldon and S. Verhuist
those problems that arise in and of itself,
but not those imposed by mythology. Dispersal and local dynamics of benthic
invertebrates,
Acknowledgements I MA. Palmer et al.
Support was provided by grant
No. 95-04-13573a of the Russian

306 TREE vol. II, no. 7 July 1996