The Quantification of Biodiversity: An Esoteric Quest or a Vital Component of

Sustainable Development?
Author(s): Thomas E. Lovejoy
Source: Philosophical Transactions: Biological Sciences , Jul. 29, 1994, Vol. 345, No. 1311,
Biodiversity: Measurement and Estimation (Jul. 29, 1994), pp. 81-87
Published by: Royal Society

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The quantification of biodiversity: an esoteric quest or a
vital component of sustainable development?
THOMAS E. LOVEJOY
Smithsonian Institution, 1000 Jefferson Drive S.W., Ste 320, Washington, D.C. 20560, U.S.A.

CONTENTS

PAGE

1. Introduction 81
2. Economic considerations 82
3. Agriculture and pest management 82
4. Pharmaceuticals 83

5. Environmental applications 83
6. Molecule-level benefits 83

7. Sustainable development and environmental management 84

8. Ecosystem function 84
References 86

SUMMARY

Biodiversity relates to sustainable development through a series of direct and indirect us

direct harvest, nature tourism, wild genes improving domestic crops, wild species contri
productivity, pest management, sources of medicine and bioremediation (biological
mental clean-up). Biodiversity relates through services, individual species indicating
change or stress, insights into the life sciences and increasingly today, through wealth g
biodiversity at the level of the molecule. Sustainable development relates to the q
biodiversity through organizing information to enable the foregoing activities. It also re
explored ways to ecosystem function, stability and resilience. Biodiversity is already a pr
environmental change in freshwater systems.

1. INTRODUCTION
and systems. This understanding is certainly sufficient
to ask whether the new-found interest in quantifying
biodiversity
Accelerating extinction rates and the Conventionis someon
esoteric quest or whether it
indeed relates
Biological Diversity have cast a new spotlight on in fundamental ways to sustainable
studies
of biodiversity and therefore, relevant
development. research
One obvious
(Smith et al. 1993). Simultaneously, the way that quantifying biodiversity
scientific
relates toin
community has developed new interests theexploring
societal imperative of sustainable devel-
opment is has
and quantifying biodiversity. Wilson (1985) that itcalled
includes straight forward biological
good housekeeping,
for a complete inventory of biological diversity thatand
is, the inventory, survey and
monitoring of
has recognized that learning how many forms of life basic biological resources. This is the
basic motivationquest.
inhabit the planet is a legitimate scientific behind the recent establishment of the
National
Similarly, Janzen (1993) has set forth theBiological
goal of Survey
an in the United States and the
general recommendation
All Taxon Biodiversity Inventory (ATBI) to quantifyemanating from the United
and describe biological diversity inNations Conference
a discrete on Environment and Development
unit
of landscape. These are certainly valid (UNCED) for thegoals
scientific institution of biological surveys in all
of great relevance to evolutionary,nations.
ecological and
To most
biogeographical science, but do they or howpeople,
do theyhowever, biological survey and
relate to the current environmental and social monitoring seems an abstruse scientific activity
imperative of sustainable development? concerned with a vast variety of weirdly named
species about which no one has heard and which are
Sustainable development is far from tightly defined
of little
and we are far from a rigorous recipe of how to relevance to human society. Unfortunately,
the majority of human society is ignorant of the extent
recognize it when we see it. Nonetheless, it is generally
to which we depend on or benefit from a great array
agreed that it must include maintenance of ecological
of other forms of life. Although such people might
systems, natural resource bases, and social structures

Phil. Trans. R. Soc. Lond. B (1994) 345, 81-87 ? 1994 The Royal Society
Printed in Great Britain 81

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82 T. E. Lovejoy Quantification of biodiversity

be willing to acknowledge that as living entities industrialized countries such as the United States
people need biological resources, most labour under (Lipske 1994).
the illusion that all that really matters is a handful of Another development in the contribution of
plant and animal species used as foods enlivened by a biodiversity to economic development in recent years
few more used as spices, with a couple of domestic has been nature tourism (Boo 1990). In certain
animals such as dogs or cats thrown in for amusement. countries like Kenya and Costa Rica, nature-oriented
tourism vies for place as the number one foreign
2. ECONOMIC CONSIDERATIONS
exchange earner. Ecotourism seems to be developing
rapidly in a large number of countries. Twenty-nine
In contrast, biological resources represent
million United
a significant
States citizens participated in a total of
contribution to economic activity and 310- million naturethey
provided trips in 1980. Of these citizens,
are managed prudently - therefore to sustainable
1031000 made 4067000 trips to foreign countries.
development. Prescott-Allen & Prescott-Allen (1986)
(U.S. Fish & Wildlife Service 1982) It is still difficult
produced the first analysis of the importance to get an accurate estimate as to the extent of world
of wild
species to the United States economy. For the ecotourism but it is certainly on the order of billions of
period 1976-1980 they estimated that 4.5% of dollars annually (Lindberg 1991).
the Gross Domestic Product (GDP) was attributable By themselves, extractive reserves and ecotourism
to wild species. Of this, 91% consisted of wild would seem to be relatively weak arguments for the
harvested resources (27 billion dollars or 4.1% importance of biodiversity and its quantification to
of GDP), 7% was of wild resources in support of sustainable development. Indeed ecotourism can often
agriculture and 2% represented wildlife-based recrea- unfairly be given an elitist cast. It is important to
tion. Their analysis of the living resource base showed note, however, that the same wild areas of value for
17% to be wild resources and a further 5% composedthose economic uses can serve as resource reserves
of new domesticates, wild genetic resources and thefor other, often very sophisticated, economic uses of
contribution of wild pollinators. In less developed biodiversity.
nations, the wild element in economic productivity
probably looms larger, although in many instances it3. AGRICULTURE AND PEST MANAGEMENT
is likely to occur unmeasured by the formal economy
(IUCN 1993). Among the uses of biodiversity for economic
A recognition of the value of wild species to which ordinarily escape mainstream econo
economic activity is inherent in the experimentationculus, is the use of genetic traits from wild r
with extractive reserves in tropical forest countries like of domestic crop species. The internationa
Brasil. In Brasil, such experimentation is largelyfor various crops such as rice or wheat are
focused on Brasil nuts and wild rubber. In addition,continually turning to wild relatives for disease and
however, the Babassu palm forests of Amazonia holdpest resistance. Similarly, that part of the forestry
considerable promise as extractive reserves (Anderson industry which is actively engaged in reforestation
et al. 1991) which is not surprising given the highand plantation forestry is continually drawing on
economic importance of the palm family in general.wild genes to improve varieties. This use of wild
In southeast Asia, the various wild species of thegenetic resources extends beyond wild relatives of crop
climbing palms known as rattans are an importantor tree species to other species of great value to
forest product. Indonesia's 1985 exports of unworkedproduction such as nitrogen-fixing bacteria or myco-
rattan was valued at $97 million. World trade in rhizzal fungi. Relatively small increases in productivity
rattan products that year was estimated at $2.7 billion
or resistance to pests and disease can yield large
(Kiew 1991 a; Mogea 1991; Pearce 1991). increments in profitability.
Prior to the establishment of tropical areas
Yet another way in which biodiversity contributes
specifically as extractive reserves, Peters et. al. (1989)
to economic activity is in integrated pest management
conducted a provocative analysis of the economic to foster the more judicious use of pesticides. For
potential of various extraction products fromexample,
Ama- two fungi, Metarhizium fiavoviride and
zonian forests in contrast to the yield from forests
Beauveria bassiana, are currently proving effective in
converted to cattle pasture. They demonstrated that
field testing against migratory locusts and agricul-
extractive activity was potentially far more lucrative,
turally problematical grasshoppers (Leary 1994). This
but what was lacking was any way to estimate theconfined to agriculture and forestry but to other
is not
market and therefore the price stability for the
areas of resource management as well. Two species of
products. A somewhat similar analysis hasOpuntia
been cactus had once spread in such density across
made of the economic value of traditional medicines
an area of Australia equivalent to the size of the
from two forest areas of Belize (Balick & MendelsohnUnited Kingdom that half that area was rendered
1992). For such schemes to become more than isolateduseless. Introduction of the moth, Cactoblastis cactorum,
examples, there is a need to develop means ofin 1925 eliminated cactus infestation from vast areas
transport of products which in turn do not attract and has controlled it subsequently (Waterhouse
destructive development. Nonetheless, non-timber 1991).
forest products already play a significant role in Similar examples of pest control occur all the time.
informal and formal economies (Plotkin & Famolare The new volumes on the weevils of Australia might
1992). This is true but probably to a lesser extent seem
in the ultimate in esoterica to some, and who, one

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 Quantification of biodiversity T. E. Lovejoy 83

might ask, has ever heard of a 'good' weevil? Yet these biodiversity will represent a pool of biologically
volumes (Zimmerman 1991) contain accounts of Lake active molecules pretested in nature which can be of
Moondara, Queensland and the river at Imbuando immense value. For some time it has been thought
Village, East Sepik, Papua New Guinea. There, that the odds of a potential lead in screening extracts
choking mats of the Brazilian waterweed Salvia from wild species for potential medical activity of one
molesta once virtually suffocated the natural produc- sort or another are but one in 10000 (McChesney
tivity and value for fish production of these fresh 1992). That coupled with the long lag times needed to
water ecosystems. Introduced Brazilian Cyrtobagous translate a lead into a commercially successful
weevils led to the virtual elimination of the exotic medicine leads to highly discounted present value
plant and subsequently, the waterbodies have recov- (Reid et al. 1993). The numbers seem to be changing
ered. In a partial echo, another weevil, Euhrychiopsis
in favour of increasing present value as screening
lecontei, once a pest of an indigenous aquatic plant,techniques improve, and particularly with simul-
mutated to a form which effectively attacks Eurasian taneous screening for multiple types of activity.
Prospecting for nature's biochemical riches (Eisner
water milfoil, Myriophyllum spicatum, which has become
a serious problem species in the United States 1990) seems to be growing in importance.
(Sheldon 1993).
5. ENVIRONMENTAL APPLICATIONS
4. PHARMACEUTICALS
The value of biodiversity to waste mana
In addition, unbeknown to most, wild species haveclean-up problems, thro
environmental
been a vitally important source of medicines nique known at least
as bioremediation, is increa
since Hippocrates was prescribing infusions The discovery
of willow of microorganisms with o
bark (the precursor of aspirin) as anlisms and In
analgesic. appetites
the can greatly facilit
United States, it is estimated that 25% suchof prescrip-One of the more intrig
problems.
tions have active ingredients with plant bacteriumorigin.
found in the sediments of the Potomac river
Sales
of these have been estimated at 4.5 billion dollars in which has the ability to break down the ozone
1980 and 15.5 billion dollars in 1990 (P. P. Principe, destroying chemicals known as chlorofluorocarbons
unpublished data). In the United States, Canada, (CFCs) (Lovley & Woodward 1992). Already there
Europe, Japan and Australia, prescription and non- are commercial applications of microbes which break
prescription drugs of plant origin were estimated to down organic molecules such as occur in oil spills. The
have a market value in 1985 of 43 billion dollars major economic potential of bioremediation may lie in
(Principe 1989). What is not included in these removing certain elements from waste streams before
estimates is the economic activity generated by a release from a factory.
healthier populace.
One of the more promising discoveries in recent6. MOLECULE-LEVEL BENEFITS
years has been taxol which was initially isolated from
the Pacific yew tree (Taxus brevifolia). It is being used I believe we are in transition to an era
in the treatment of ovarian and breast cancers. The benefits will accrue to human society an
vine, Ancistrocladus korupensis, first collected development
from from wild resources becau
Korup National Park in Cameroon, yielded the generated at the level of the molecule
compound michellamine B, so successful in protecting 1992). A new bioflocculant derived from
human cells from the HIV virus in vitro that it is the potatorum, a tree in Andrha Pradesh, a
first natural compound to move to animal testing for able to clean up uranium and other long-lived
AIDS research (Washington Post 1994; Stix 1993). isotopes of nuclear wastes. It appears to have similar
Squalamine, an aminosterol from the dogfish (Squalus ability to remove heavy metals such as cadmium
acanthias), displays antiparasitic, antifungal, anti- and mercury from effluent (Jayaraman 1993).
protozoan and antibacterial activity, including Azadirachtin from the neem tree (Azadirachta indica)
against Streptococcus, in vitro (Moore et al. 1993). An from India has recently become a commercially
Ecuadorian arrow poison frog, Epipedobates tricolor,available insecticide (Stone 1993). Marine organisms
secretes a compound which blocks pain 200 times as contain substances under investigation for various
effectively as morphine but is not an opioid (Daly uses: discodermolide from a Bahamian sponge,
1994; Spande et al. 1992). Discodermia sp., may be useful in suppressing organ
Very often production of a pharmaceutical product rejection after transplant operations; macrolactins
initially requires a lot of material harvested in nature from the Bahamas sea floor mud bacteria appear
to extract the active ingredient but this is often promising in inhibiting growth of melanoma and
superseded by the ability to synthesize. Although this colon cancer cells as well as the herpes simplex and
means that in one sense the biological source is no HIV (Fenical 1989, 1993).
longer necessary, it is important to recognize that it Genetic engineering now makes it possible to
derives from the original inspiration, that is, the introduce desirable genetic traits from one species
template provided by a wild species. into another which is not closely related. Pest resistant
While it is possible the day may come when genes from Bacillus thuringiensis have been transferred
computers can be used to design molecules to affectto a variety of crop species (Gasser & Fraley 1992). A
particular disease organisms, even at such a timefreeze-resistant strain of tobacco has been produced

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84 T. E. Lovejoy Quantification of biodiversity

by inclusion of a gene from winter flounder (Gladwell Vaccinia (cow pox) virus and by antibiotics and
1990). The development of the 'Flavr Savr' tomato Penicillium mould. Humanity's stake in being able to
about which there has been so much controversy in understand evolution is presumably more than a
the United States actually involves only the manipu- trivial and esoteric matter.
lation of tomato genes to delay the softening which The properties that collectively make each species a
normally comes with ripening so that ripe tomatoes vast store of information (Wilson 1985) also make
can be shipped long distances without rapid spoiling them the most sensitive indicators of environmental
(Singer 1993). There has been considerable trepida- change. The concentration of chlorinated hydro-
tion about genetically engineered organisms among carbons as these molecules passed up foodchains
the public. Once it is better understood that, like most culminated in drastic declines of top predators such
technologies, genetic engineering is essentially neutral as peregrine falcons (Falco peregrinus) making them
and that benefits or problems relate instead to how the first indicators of the 'Silent Spring' problems
the technology is applied, there should be rapid predicted by Rachel Carson. The toxic effects on non-
economic return generated from biodiversity in this target species such as the European oyster (Ostrea
fashion. edulis) led to re-evaluation of the use and regulation of
Probably, the most significant example of the power tributyl tin as an antifouling agent for nautical paints
of a single molecule from the wild to generate new and coatings. Unhealthy and dying red spruce (Picea
possibilities and economic development involves the rubens) were the first visible indicator of the acid rain
polymerase chain reaction for the discovery of which problem in North America.
Kary B. Mullis shared the 1993 Nobel Prize in
chemistry. This reaction, which permits multipli- 8. ECOSYSTEM FUNCTION
cation of DNA billions of times in a few hours, is
now fundamental to diagnostic medicine as well asAnother argument often advanced about
much of biotechnology. It depends on a heat-resistant biodiversity concerns the public services pe
enzyme discovered by Thomas Brock in the bacterium the natural aggregations of biodiversity
Thermus aquaticus which occurs naturally in hot springs ecosystems. There is no question that th
in Yellowstone National Park (Mullis 1990). metabolism of the planet is conducted by t
The reaction, which is so basic in molecular biology biodiversity and biomass, or that ecosy
as to be referred to generally as PCR, relates to important roles in the cycling of nutrie
probably the most important and least recognized energy and biogeochemical cycling in gen
relationship between biodiversity and sustainable it comes down to specifics of the relationsh
development, namely the value of wild species as biodiversity and the functioning of ecos
intellectual resources. Biodiversity in essence repre-picture is far more complex and only f
sents the most fundamental library in support of the understood (Schulze & Mooney 1993). Fo
life sciences: tens of millions of species with unique setswatershed regulation might seem provided
of properties, processes and 'solutions' to unique sets ofa plantation forest as by a highly diverse n
biological histories and challenges. These are not, of Agricultural ecosystems drastically sim
course the only 'solutions', they are the ones whichthe pragmatic purposes of production, sugg
have evolved and survived to this moment. Many a modest link between biodiversity and
others have occurred in the past and the number function. Five different treatments in Costa Rica
which might evolve in the future is inestimable. Yet(bare ground, maize monoculture, natural succession
those currently existing tell us interesting things aboutmimicked succession and enriched succession) indicate
how evolution has worked and can work. The increases in soil nutrient pools for the various
discovery of the hydrothermal vent communities successional treatments
in relative to the monoculture
the 1970s, which demonstrated that entire or commu-
the bare ground (Ewel et al. 1991). This certainl
nities could depend on the primal energy of suggests
the Eartha positive relation between ecosystem func
rather than solar radiation and that organisms tion and biodiversity, although results might hav
could
exist at temperatures greater than the boiling differed
pointif ofother crop species had been used for
water, demonstrate just how far we are from the under-
monoculture. Similarly, low number multiple
standing even the most basic dimensions of lifesystems
cropping on tend to produce greater yields than
Earth. monoculture (Swift & Anderson 1993).
In the more complex reality of most natural
7. SUSTAINABLE DEVELOPMENT AND
ecosystems, the picture is much less obvious. There
ENVIRONMENTAL MANAGEMENT are, however, some rather simple situations such as
microbial communities in Antarctica where species
The notion that biologically based development number is positively related to energy flow because
constitutes an important segment of sustainable of the greater number of chemical sources which
development because of the ability of biological can provide energy through microbial oxidation
resources to renew themselves, supports the impor- (Woodward 1993). This is analogous to the number
tance of the life sciences and the potential that can be of functional groups of microbes and other organisms
derived from examining things that occur in nature. in much more complex and rich ecosystems.
The power of concepts derived from observing nature Clearly some species play more significant roles in
to benefit society is exemplified by vaccination and ecosystem functioning than others (Vitousek & Hooper

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 Quantification of biodiversity T. E. Lovejoy 85

1993). This evolves from the concept of keystone ecosystem structure it may in fact be playing a
species (Paine 1966). Despite the identification of a significant role.
number of types of keystone species (Lawton & Perhaps more important is that the present role of
Brown 1993), the types relate primarily to ecosystem a rare species may provide little hint of its role or
structure and it does not follow that a keystone species usefulness at other times when environmental condi-
is necessarily vital for ecosystem function. In contrast, tions vary (Main 1982; Lovejoy 1988). Gap species in
however, fungal transfer of nutrients by mycorhizas a tropical forest are an example, as are the many
can be significant. species which require fire to enhance germination.
An interesting dimension of the concept of keystone Ehrlich (1993) points out that it would be difficult to
species is provided by the Chesapeake Bay ecosystem divine the importance of Cactoblastis cactorum with
(Lovejoy 1993). This ecosystem is of considerable respect to the Australia Opuntia story from its rather
extent, draining significant portions of New York, rare and infrequent occurrence in those ecosystems
Pennsylvania, Maryland and Virginia and includes today. A particularly interesting example is a yeast with
a vast estuary of 103 040 km2. The American oyster the ability to reduce mercury (Brunner & Bott 1974).
(Crassostrea virginica) is today estimated to filter a Normally rare in aquatic communities, it becomes
volume of water equal to the entire bay once a year, abundant during episodes of mercury contamination
but prior to the drastic decline of its population (natural or anthropogenic), cleans mercury out of the
filtered that same volume once a week (Newell 1988). ecosystem and perforce becomes rare once again.
Although oysters never occurred throughout the The actual quantification of biodiversity at the level
entire bay, their effect on water clarity and thus the of the ecosystem has already proven to be of consider-
general ecology must have been considerable. able pragmatic value. Working on freshwater systems,
The relatively few experimental manipulations of Patrick and others have developed a considerable body
ecosystems shed a little light on the relationship of work relating biodiversity to ecosystem stress (Patrick
between biodiversity and ecosystem function. For 1949, 1953, 1961, 1968; Patrick et al. 1968, 1969).
example, the recovery of ecosystem function by the Essentially the numbers and kinds of species as well as
deforested Hubbard Brook watershed probably related the distribution of abundance between species provide
more to biomass than species number (Likens 1985). an immensely practical measure of the extent to which
The relationship of the stability of an ecosystem to the ecology of a river or other freshwater body has
biological diversity is a complicated one (MacArthur departed from normal. This property of quantified
1955; May 1974) but vital to any consideration of biodiversity is essentially the integral of the indicator
sustainable development. An interesting aspect is property of individual species as enhanced or modified
resistance to invasion by alien species, a serious by ecosystem structure and internal interactions of those
problem in the United States (U.S. Congress, Office systems. Karr (1991) has used this approach to develop
of Technology Assessment 1993) and elsewhere. There an Index of Biotic Integrity (IBI).
does appear to be a greater resistance to invasion by This principle is transferrable, at least in part to
more species rich ecosystems, a concept deriving terrestrial ecosystems. An experimental grass plot
from pioneering ecologist Charles Elton (1958). Most subject to application of nitrogen fertilizer at
examples of efforts of alien species relate to the effect Rothamsted showed reduction in species number
on indigenous species; but one relating to ecosystem and decreasing equitability of abundance among
function involves the invasion of Myrica faya in species over decades. The reverse was true of an
a Hawaii lava field ecosystem. This led to a tenfold abandoned agricultural field in Illinois over a 40 year
change in nitrogen fixation. The recent work of period (May 1978). The biodiversity of terrestrial
Tilman & Downing (1994) show prairie plots with ecosystems may in some instances reflect the extent of
greater plant diversity are more resistant to severe an ability to resist stress. A set of experiments involving
drought than plots with lower plant diversity. nutrient enrichment of old fields in New York resulted
The relationship of biodiversity and ecosystem in the more diverse plant community being less
function raises the little-explored topic of the extent sensitive to the treatment. Interestingly, this did not
to which there is redundancy (Walker 1992; Lawton occur at the next trophic level (McNaughton 1993).
& Brown 1993) of species' function in ecosystems. This relationship between biodiversity and eco-
Undoubtedly, there are species which perform rather system stress suggests that a measure of biodiversity
similar roles (Hutchinson 1961) although with furthersuch as that developed by Patrick for freshwater
knowledge they may be distinguishable (Steinberg & ecosystems could be very useful in the rapidly
Geller 1993). Redundancy can, in any case, be useful emerging field of ecosystem management. Simple
in living systems. measures such as alpha (Fisher et al. 1943) should
Such analysis can be taken a step further to inquire be explored. This should not be to the exclusion of
whether rare species play important roles in eco- particular indicator species or taxa which can be of
systems and their function. In highly diverse tropical extreme utility in particular ecosystems and situations.
ecosystems such as those of tropical forests or coral In Australia, for example, ant diversity has proven to
reefs, there is such a quantity of rare species that taken be of great value for monitoring ecosystem recovery
collectively they do play an important role in (Majer 1983). The utility of these kinds of approaches
ecosystem function. In addition, a rare species may to monitoring the state of ecosystems will depend in
be rare because it is both large and at the top of the turn on the progress made in describing, studying and
foodchain where at least from the perspective of quantifying life on Earth.

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86 T. E. Lovejoy Quantification of biodiversity

I am grateful to the following for helpful discussions and
input to this contribution: Ann Bartuska, Sarah A. Boren,
John W. Daly, Paul Dresler, Ryan H. Dwight, William
Fenical, John H. Gibbons, Frances C. James, Daniel H.
Janzen, Peter R. Jutro, Robert M. May, Ebbe S. Nielsen,
Robin I. O'Malley, Ruth Patrick, Richard B. Primack and
Sallie Sheldon.
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