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Issue 2
Issues in Ecology is an official publication of the Ecological Society of America, the nations
leading professional society of ecologists. Founded in 1915, ESA seeks to promote the
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For more information, contact the Ecological Society of America, 2010 Massachusetts Av-
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Published by the Ecological Society of America Number 2, Spring 1997
Issues in Ecology by Natural Ecosystems
Supplied to Human Societies
Ecosystem Services: Benefits
Issues in Ecology Number 2 Spring 1997
Ecosystem Services:
Benefits Supplied to Human
Societies by Natural Ecosystems
SUMMARY
Human societies derive many essential goods from natural ecosystems, including seafood, game animals, fodder,
fuelwood, timber, and pharmaceutical products. These goods represent important and familiar parts of the economy.
What has been less appreciated until recently is that natural ecosystems also perform fundamental life-support services
without which human civilizations would cease to thrive. These include the purification of air and water, detoxification
and decomposition of wastes, regulation of climate, regeneration of soil fertility, and production and maintenance of
biodiversity, from which key ingredients of our agricultural, pharmaceutical, and industrial enterprises are derived. This
array of services is generated by a complex interplay of natural cycles powered by solar energy and operating across a
wide range of space and time scales. The process of waste disposal, for example, involves the life cycles of bacteria as well
as the planet-wide cycles of major chemical elements such as carbon and nitrogen. Such processes are worth many
trillions of dollars annually. Yet because most of these benefits are not traded in economic markets, they carry no price
tags that could alert society to changes in their supply or deterioration of underlying ecological systems that generate
them. Because threats to these systems are increasing, there is a critical need for identification and monitoring of
ecosystem services both locally and globally, and for the incorporation of their value into decision-making processes.
Historically, the nature and value of Earths life support systems have largely been ignored until their disruption or
loss highlighted their importance. For example, deforestation has belatedly revealed the critical role forests serve in
regulating the water cycle -- in particular, in mitigating floods, droughts, the erosive forces of wind and rain, and silting
of dams and irrigation canals. Today, escalating impacts of human activities on forests, wetlands, and other natural
ecosystems imperil the delivery of such services. The primary threats are land use changes that cause losses in biodiversity
as well as disruption of carbon, nitrogen, and other biogeochemical cycles; human-caused invasions of exotic species;
releases of toxic substances; possible rapid climate change; and depletion of stratospheric ozone.
• Many of the human activities that modify or destroy natural ecosystems may cause deterioration of ecological
services whose value, in the long term, dwarfs the short-term economic benefits society gains from those activities.
• Considered globally, very large numbers of species and populations are required to sustain ecosystem services.
• The functioning of many ecosystems could be restored if appropriate actions were taken in time.
We believe that land use and development policies should strive to achieve a balance between sustaining vital
ecosystem services and pursuing the worthy short-term goals of economic development.
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Issues in Ecology Number 2 Spring 1997
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Issues in Ecology Number 2 Spring 1997
generating those ecosystem goods that are traded in the directly for food, drink, spices, fiber, timber, pharmaceu-
marketplace. As a result, this lack of awareness helps ticals, and industrial products such as waxes, rubber, and
drive the conversion of natural ecosystems to oils. Even if one were highly selective, the list could amount
human-dominated systems (e.g., wheatlands or oil palm to hundreds or even thousands of species. And that would
fields), whose economic value can be expressed, at least only be a start, since one would then need to consider
in part, in standard currency. The second reason to focus which species are crucial to supporting those used di-
on natural ecosystems is that many human-initiated dis- rectly: the bacteria, fungi, and invertebrates that help
ruptions of these systems -- such as introductions of ex- make soil fertile and break down wastes and organic
otic species, extinctions of native species, and alteration matter; the insects, bats, and birds that pollinate flow-
of the gaseous composition of the atmosphere through ers; and the grasses, herbs, and trees that hold soil in
fossil fuel burning -- are difficult or impossible to reverse place, regulate the water cycle, and supply food for ani-
on any time scale relevant to society. Third, if awareness mals. The clear message of this exercise is that no one
is not increased and current trends continue, humanity knows which combinations of species -- or even approxi-
will dramatically alter Earths remaining natural ecosys- mately how many -- are required to sustain human life.
tems within a few decades (Daily 1997a, b). Rather than selecting species directly, one might
The lack of attention to the vital role of natural try another approach: Listing the ecosystem services
ecosystem services is easy to understand. Humanity came needed by a lunar colony and then guessing at the types
into being after most ecosystem services had been in and numbers of species required to perform each. Yet
operation for hundreds of millions to billions of years. determining which species are critical to the functioning
These services are so fundamental to life that they are of a particular ecosystem service is no simple task. Let
easy to take for granted, and so large in scale that it is us take soil fertility as an example. Soil organisms are
hard to imagine that human activities could irreparably crucial to the chemical conversion and physical transfer
disrupt them. Perhaps a thought experiment that re- of essential nutrients to higher plants. But the abun-
moves these services from the familiar backdrop of the dance of soil organisms is absolutely staggering. Under
Earth is the best way to illustrate both the importance a square-yard of pasture in Denmark, for instance, the
and complexity of ecosystem services, as well as how soil is inhabited by roughly 50,000 small earthworms
ill-equipped humans are to recreate them. Imagine, for and their relatives, 50,000 insects and mites, and nearly
example, human beings trying to colonize the moon. 12 million roundworms. And that tally is only the begin-
Assume for the sake of argument that the moon had ning. The number of soil animals is tiny compared to the
already miraculously acquired some of the basic condi- number of soil microorganisms: a pinch of fertile soil may
tions for supporting human life, such as an atmosphere, contain over 30,000 protozoa, 50,000 algae, 400,000
a climate, and a physical soil structure similar to those fungi, and billions of individual bacteria
on Earth. The big question facing human colonists would (Overgaard-Nielsen 1955; Rouatt and Katznelson 1961;
then be, which of Earths millions of species would need Chanway 1993). Which must colonists bring to the moon
to be transported to the moon to make that sterile sur- to assure lush and continuing plant growth, soil renewal,
face habitable? waste disposal, and so on? Most of these soil-dwelling
One could tackle that question systematically species have never been subjected to even cursory in-
by first choosing from among all the species exploited spection: no human eye has ever blinked at them through
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Issues in Ecology Number 2 Spring 1997
a microscope, no human hand has ever typed out a name erated by sport fishing activities, it raises the total to
or description of them, and most human minds have never $46 billion (Felder and Nickum 1992, cited in Postel and
spent a moment reflecting on them. Yet the sobering Carpenter 1997). The future of these fisheries is in ques-
fact is, as E. O. Wilson put it: they dont need us, but we tion, however, because fish harvests have approached or
need them (Wilson 1987). exceeded sustainable levels virtually everywhere. Nine
of the worlds major marine fishing areas are in decline
THE CHARACTER OF ECOSYSTEM SERVICES due to overfishing, pollution, and habitat destruction.
(UNFAO 1993; Kaufman and Dayton 1997).
Moving our attention from the moon back to Earth, let Turning our attention to the land, grasslands are
us look more closely at the services nature performs on an important source of marketable goods, including ani-
the only planet we know that is habitable. Ecosystem mals used for labor (horses, mules, asses, camels, bul-
services and the systems that supply them are so inter- locks, etc.) and those whose parts or products are con-
connected that any classification of them is necessarily sumed (as meat, milk, wool, and leather). Grasslands
rather arbitrary. Here we briefly explore a suite of were also important as the original source habitat for
overarching services that operate in ecosystems world- most domestic animals such as cattle, goats, sheep, and
wide. horses, as well as many crops, such as wheat, barley,
rye, oats, and other grasses (Sala and Paruelo 1997). In
Production of Ecosystem Goods a wide variety of terrestrial habitats, people hunt game
Humanity obtains from natural ecosystems an animals such as waterfowl, deer, moose, elk, fox, boar
array of ecosystem goods organisms and their parts and other wild pigs, rabbits, and even snakes and mon-
and products that grow in the wild and that are used keys. In many countries, game meat forms an important
directly for human benefit. Many of these, such as fishes part of local diets and, in many places, hunting is an
and animal products, are commonly traded in economic economically and culturally important sport.
markets. The annual world fish catch, for example, Natural ecosystems also produce vegetation used directly
amounts to about 100 million metric tons and is valued by humans as food, timber, fuelwood, fiber, pharmaceu-
at between $50 billion and $100 billion; it is the leading ticals and industrial products. Fruits, nuts, mushrooms,
source of animal protein, with over 20% of the popula- honey, other foods, and spices are extracted from many
tion in Africa and Asia dependent on fish as their pri- forest species. Wood and other plant materials are used
mary source of protein (UNFAO 1993). The commercial in the construction of homes and other buildings, as well
harvest of freshwater fish worldwide in 1990 totaled as for the manufacture of furniture, farming implements,
approximately 14 million tons and was valued at about paper, cloth, thatching, rope, and so on. About 15 per-
$8.2 billion (UNFAO 1994). Interestingly, the value of cent of the worlds energy consumption is supplied by
the freshwater sport fishery in the U.S. alone greatly fuelwood and other plant material; in developing coun-
exceeds that of the global commercial harvest, with di- tries, such biomass supplies nearly 40 percent of en-
rect expenditures in 1991 totaling about $16 billion. ergy consumption (Hall et al. 1993), although the por-
When this is added to the value of the employment gen- tion of this derived from natural rather than
oats.
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Issues in Ecology Number 2 Spring 1997
human-dominated ecosystems is undocumented. In addi- As described in the previous section, biodiversity
tion, natural products extracted from many hundreds of is a direct source of ecosystem goods. It also supplies
species contribute diverse inputs to industry: gums and the genetic and biochemical resources that underpin our
exudates, essential oils and flavorings, resins and oleo- current agricultural and pharmaceutical enterprises and
resins, dyes, tannins, vegetable fats and waxes, insecti- may allow us to adapt these vital enterprises to global
cides, and multitudes of other compounds (Myers 1983; change. Our ability to increase crop productivity in the
Leung and Foster 1996). The availability of most of face of new pests, diseases, and other stresses has de-
these natural products is in decline due to ongoing habi- pended heavily upon the transfer to our crops of genes
tat conversion. from wild crop relatives that confer resistance to these
challenges. Such extractions from biodiversitys genetic
Generation and Maintenance of Biodiversity library account for annual increases in crop productiv-
Biological diversity, or biodiversity for short, re- ity of about 1 percent, currently valued at $1 billion (NRC
fers to the variety of life forms at all levels of organiza- 1992). Biotechnology now makes possible even greater
tion, from the molecular to the land- use of this natural storehouse of ge-
scape level. Biodiversity is generated netic diversity via the transfer to crops
and maintained in natural ecosystems, of genes from any kind of organism
where organisms encounter a wide not simply crop relativesand it
variety of living conditions and chance promises to play a major role in fu-
events that shape their evolution in ture yield increases. By the turn of
unique ways. Out of convenience or the century, farm-level sales of the
necessity, biodiversity is usually quan- products of agricultural biotechnol-
tified in terms of numbers of species, ogy, just now entering the market-
and this perspective has greatly influ- place, are expected to reach at least
enced conservation goals. It is im- $10 billion per year (World Bank
portant to remember, however, that 1991, cited in Reid et al. 1996).
the benefits that biodiversity supplies In addition to sustaining the produc-
to humanity are delivered through tion of conventional crops, the
populations of species residing in liv- biodiversity in natural ecosystems may
ing communities within specific physi- include many potential new foods.
Photo by Taylor Ricketts
cal settings in other words, through Human beings have utilized around
complex ecological systems, or eco- 7,000 plant species for food over the
systems (Daily and Ehrlich 1995). For course of history and another 70,000
human beings to realize most of the plants are known to have edible parts
aesthetic, spiritual, and economic ben- (Wilson 1989). Only about 150 food
efits of biodiversity, natural ecosys- Figure 4-Harpoon-whaling in Flores, In- plants have ever been cultivated on a
tems must therefore be accessible. The donesia. The oceans are a key source large scale, however. Currently, 82
continued existence of coniferous tree of animal protein for the human popu- plant species contribute 90 percent
species somewhere in the world would lation. of national per-capita supplies of food
not help the inhabitants of a town in- plants (Prescott-Allen and
undated by flooding because of the clearing of a pine Prescott-Allen 1990), although a much smaller number
forest upstream. Generally, the flow of ecosystem goods of these supply the bulk of the calories humans consume.
and services in a region is determined by the type, spa- Many other species, however, appear more nutritious or
tial layout, extent, and proximity of the ecosystems sup- better suited to the growing conditions that prevail in
plying them. Because of this, the preservation of only important regions than the standard crops that domi-
one minimum viable population of each non-human spe- nate world food supply today. Because of increasing
cies on Earth in zoos, botanical gardens, and the worlds salinization of irrigated croplands and the potential for
legally protected areas would not sustain life as we know rapid climate change, for instance, future food security
it. Indeed, such a strategy, taken to extreme, would lead may come to depend on drought- and salt-tolerant vari-
to collapse of the biosphere, along with its life support eties that now play comparatively minor roles in agricul-
services. ture.
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Issues in Ecology Number 2 Spring 1997
Turning to medicinal resources, a recent survey age 20,000 years ago, for example, much of Europe and
showed that of the top 150 prescription drugs used in North America were covered by mile-thick ice sheets.
the United States, 118 are based on natural sources: While the global climate has been relatively stable since
74% on plants, 18% on fungi, 5% on bacteria, and 3% the invention of agriculture around 10,000 years ago,
on one vertebrate (snake) species. Nine of the top ten periodic shifts in climate have affected human activities
drugs in this list are based on natural plant products and settlement patterns. Even relatively recently, from
(Grifo and Rosenthal, in press, as cited in Dobson 1995). 1550-1850, Europe was significantly cooler during a
The commercial value of pharmaceuticals in the devel- period known as the Little Ice Age. Many of these changes
oped nations exceeds $40 billion per year (Principe 1989). in climate are thought to be caused by alterations in
Looking at the global picture, approximately 80% of the Earths orbital rotation or in the energy output of the
human population relies on traditional medical systems, sun, or even by events on the Earth itselfsudden per-
and about 85% of traditional medicine involves the use turbations such as violent volcanic eruptions and aster-
of plant extracts (Farnsworth et al. 1985). oid impacts or more gradual tectonic events such as the
Saving only a single popula- uplift of the Himalayas. Remarkably,
tion of each species could have an- climate has been buffered enough
other cost. Different populations of through all these changes to sustain
the same species may produce differ- life for at least 3.5 billion years
ent types or quantities of defensive (Schneider and Londer 1984). And
chemicals that have potential use as life itself has played a role in this buff-
pharmaceuticals or pesticides ering.
(McCormick et al. 1993); and they Climate, of course, plays a
may exhibit different tolerances to major role in the evolution and distri-
environmental stresses such as bution of life over the planet. Yet most
drought or soil salinity. For example, scientists would agree that life itself
the development of penicillin as a is a principal factor in the regulation
therapeutic antibiotic took a full 15 of global climate, helping to offset the
years after Alexander Flemings fa- effects of episodic climate oscillations
Photo by Paul R. Ehrlich
to potentially large increases in surface runoff, along with Services Supplied by Soil
nutrient and soil loss. A classic example comes from the Soil represents an important component of a
experimental clearing of a New Hampshire forest, where nations assets, one that takes hundreds to hundreds of
herbicide was applied to prevent regrowth for a 3-year thousands of years to build up and yet very few years to
period after the clearing. The result was a 40 percent be lost. Some civilizations have drawn great strength
increase in average stream flow. During one four-month from fertile soil; conversely, the loss of productivity
period of the experiment, runoff was more than 5 times through mismanagement is thought to have ushered many
greater than before the clearing (Bormann 1968). On a once flourishing societies to their ruin (Adams 1981).
much larger scale, extensive deforestation in the Hima- Today, soil degradation induced by human activities af-
layan highlands appears to have exacerbated recent flood- flicts nearly 20 percent of the Earths vegetated land
ing in Bangladesh, although the relative roles of human surface (Oldeman et al. 1990).
and natural forces remain debatable (Ives and Messerli In addition to moderating the water cycle, as described
1989). In addition, some regions of the world, such as above, soil provides five other interrelated services (Daily
parts of Africa, are experiencing an increased frequency et al. 1997). First, soil shelters seeds and provides physi-
and severity of drought, possibly associated with exten- cal support as they sprout and mature into adult plants.
sive deforestation. The cost of packaging and storing seeds and of anchor-
Wetlands are particularly well-known for their role ing plant roots would be enormous without soil.
in flood control and can often reduce the need to con- Human-engineered hydroponic systems can grow plants
struct flood control structures. Floodplain forests and in the absence of soil, and their cost provides a lower
high salt marshes, for example, slow the flow of floodwa- bound to help assess the value of this service. The costs
ters and allow sediments to be deposited within the flood- of physical support trays and stands used in such opera-
plain rather than washed into downstream bays or oceans. tions total about US$55,000 per hectare (for the Nutri-
In addition, isolated wetlands such as prairie potholes in ent Film Technique Systems; FAO 1990).
the Midwest and cypress ponds in the Southeast, serve Second, soil retains and delivers nutrients to
as detention areas during times of high rainfall, delaying plants. Tiny soil particles (less than 2 microns in diam-
saturation of upland soils and overland flows into rivers eter), which are primarily bits of humus and clays, carry
and thereby damping peak flows. Retaining the integrity a surface electrical charge that is generally negative. This
of these wetlands by leaving vegetation, soils, and natu- property holds positively charged nutrientscations such
ral water regimes intact can reduce the severity and du- as calcium and magnesiumnear the surface, in prox-
ration of flooding along rivers (Ewel 1997). A relatively imity to plant roots, allowing them to be taken up gradu-
small area of retained wetland, for example, could have ally. Otherwise, these nutrients would quickly be leached
largely prevented the severe flooding along the Missis- away. Soil also acts as a buffer in the application of
sippi River in 1993. fertilizers, holding onto the fertilizer ions until they are
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Issues in Ecology Number 2 Spring 1997
required by plants. Hydroponic systems supply water recycling of nutrientsthe fourth service soils provide
and nutrients to plants without need of soil, but the mar- are two aspects of the same process. The fertility of
gin for error is much smallereven small excesses of soilsthat is, their ability to supply nutrients to plants
nutrients applied hydroponically can be lethal to plants. is largely the result of the activities of diverse species of
Indeed, it is a complex undertaking to regulate the nutri- bacteria, fungi, algae, crustacea, mites, termites, spring-
ent concentrations, pH, and salinity of the nutrient solu- tails, millipedes, and worms, all of which, as groups, play
tion in hydroponic systems, as well as the air and solu- important roles. Some bacteria are responsible for fix-
tion temperature, humidity, light, pests, and plant dis- ing nitrogen, a key element in proteins, by drawing it
eases. Worldwide, the area under hydroponic culture is out of the atmosphere and converting it to forms usable
only a few thousand hectares and is unlikely to grow by plants and, ultimately, human beings and other ani-
significantly in the foreseeable future; by contrast, glo- mals. Certain types of fungi play extremely important
bal cropped area is about 1.4 billion hectares (USDA roles in supplying nutrients to many kinds of trees. Earth-
1993). worms and ants act as mechani-
Third, soil plays a central cal blenders, breaking up and mix-
role in the decomposition of dead ing plant and microbial material and
organic matter and wastes, and this other matter (Jenny 1980). For
decomposition process also renders example, as much as 10 metric
harmless many potential human tonnes of material may pass
pathogens. People generate a tre- through the bodies of earthworms
mendous amount of waste, includ- on a hectare of land each year, re-
ing household garbage, industrial sulting in nutrient rich casts that
Figure 12-The goods and services supplied by this badly deforested and eroded region
of Madagascar are all but gone and would be difficult to restore.
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Issues in Ecology Number 2 Spring 1997
operation of this model • To what extent have vari-
earth, it proved impossible ous ecosystem services al-
to supply the material and ready been impaired? And
physical needs of the eight how are impairment and risk
Biospherians for the in- of future impairment distrib-
tended 2 years. Many un- uted in various regions of
pleasant and unexpected the globe?
problems arose, including a • How interdependent are
drop in atmospheric oxygen different ecosystem ser-
concentration to 14% (the vices? How does exploiting
level normally found at an or damaging one influence
elevation of 17,500 feet), the functioning of others?
high spikes in carbon diox- • To what extent, and over
ide concentrations, nitrous what time scale, are ecosys-
oxide concentrations high tem services amenable to
enough to impair the brain, repair or restoration?
an extremely high level of • How effectively, and at
extinctions (including 19 of how large a scale, can ex-
25 vertebrate species and all isting or foreseeable human
pollinators brought into the technologies substitute for
enclosure, which would have ecosystem services? What
ensured the eventual extinc- would be the side effects of
tion of most of the plant such substitutions?
species as well), overgrowth • Given the current state
of aggressive vines and al- Figure 13-The production of this meal benefited from many of technology and the scale
gal mats, and population ecosystem services, including natural pest control, pollina- of the human enterprise,
explosions of crazy ants, tion, maintenance of soil fertility, purification of water, and what proportion and spatial
cockroaches, and katydids. moderation of climate. pattern of land must remain
Even heroic personal efforts relatively undisturbed, lo-
on the part of the Biospherians did not suffice to make cally, regionally, and globally, to sustain the delivery
the system viable and sustainable for either humans or of essential ecosystem services?
many nonhuman species (Cohen and Tilman 1996).
CONCLUSIONS
MAJOR UNCERTAINTIES
The human economy depends upon the services performed
Society would clearly profit by further investigation into for free by ecosystems. The ecosystem services sup-
some of the following broad research questions so that plied annually are worth many trillions of dollars. Eco-
we might avoid on Biosphere 1, the earth, unpleasant nomic development that destroys habitats and impairs
surprises like those that plagued the Biosphere 2 project services can create costs to humanity over the long term
(Holdren 1991; Cohen and Tilman 1996; Daily 1997b): that may greatly exceed the short-term economic ben-
efits of the development. These costs are generally hid-
• What is the relative impact of various human activi- den from traditional economic accounting, but are none-
ties upon the supply of ecosystem services? theless real and are usually borne by society at large.
• What is the relationship between the condition of an Tragically, a short-term focus in land-use decisions often
ecosystemthat is, relatively pristine or heavily sets in motion potentially great costs to be borne by
modifiedand the quantity and quality of ecosys- future generations. This suggests a need for policies
tem services it supplies? that achieve a balance between sustaining ecosystem
• To what extent do ecosystem services depend upon services and pursuing the worthy short-term goals of
biodiversity at all levels, from genes to species to economic development.
landscapes?
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Issues in Ecology Number 2 Spring 1997
Ehrlich, P.R. and A.H. Ehrlich. 1981. Extinction. Ballantine, New
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Ehrlich, P.R., A.H. Ehrlich, and J.P. Holdren. 1977. Ecoscience: Popu-
We thank the Packard Foundation and the Pew lation, Resources, Environment. Freeman and Co., San Fran-
Foundation for financial support. cisco.
Ewel, K. 1997. Water quality improvement: evaluation of an ecosys-
tem service. Pages 329-344 in G. Daily, editor. Natures
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bia Univ. Press, New York.
expertise in this area. This report underwent peer review
Rouatt, J. and H. Katznelson.1961. A study of bacteria on the root and was approved by the Board of Editors of Issues in
surface and in the rhizosphere soil of crop plants. J. Ap- Ecology. The affiliations of the members of the panel of
plied Bacteriology 24: 164-171. scientists are:
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Dr. Gretchen C. Daily, Panel Chair, Department of Bio- environment. All reports undergo peer review and must
logical Sciences, Stanford University, Stanford, CA 94305 be approved by the editorial board before publication.
Dr. Susan Alexander, Earth Systems Science and Policy, Editorial Board of Issues in Ecology
California State University, Monterey Bay, 100 Campus Dr. David Tilman, Editor-in-Chief, Department of Ecol-
Center, Seaside, CA 93955 ogy, Evolution and Behavior, University of Minnesota,
St. Paul, MN 55108-6097. E-mail: tilman@lter.umn.edu
Dr. Paul R. Ehrlich, Department of Biological Sciences,
Stanford University, Stanford, CA 94305 Board members
Dr. Stephen Carpenter, Center for Limnology, University
Dr. Larry Goulder, Department of Economics, Stanford of Wisconsin, Madison, WI 53706
University, Stanford, CA 94305 Dr. Deborah Jensen, The Nature Conservancy, 1815
North Lynn Street, Arlington, VA 22209
Dr. Jane Lubchenco, Department of Zoology, Oregon Dr. Simon Levin, Department of Ecology & Evolutionary
State University, Corvallis, OR 97331 Biology, Princeton University, Princeton, NJ
08544-1003
Dr. Pamela A. Matson, Environmental Science Policy and Dr. Jane Lubchenco, Department of Zoology, Oregon
Management, University of California, Berkeley, CA State University, Corvallis, OR 97331-2914
94720 Dr. Judy L. Meyer, Institute of Ecology, The University of
Georgia, Athens, GA 30602-2202
Dr. Harold A. Mooney, Department of Biological Sciences, Dr. Gordon Orians, Department of Zoology, University
Stanford University, Stanford, CA 94305 of Washington, Seattle, WA 98195
Dr. Lou Pitelka, Appalachian Environmental Laboratory,
Dr. Sandra Postel, Global Water Policy Project, 107 Lark- Gunter Hall, Frostburg, MD 21532
spur Drive, Amherst, MA 01002 Dr. William Schlesinger, Departments of Botany and
Geology, Duke University, Durham, NC 27708-
Dr. Stephen H. Schneider, Department of Biological Sci- 0340
ences, Stanford University, Stanford, CA 94305
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About the Science Writer Special thanks to the U.S. Environmental Protec-
Yvonne Baskin, a science writer, edited the re- tion Agency Office of Sustainable Ecosystems and Com-
port of the panel of scientists to allow it to more effec- munities for supporting printing and distribution of this
tively communicate its findings with non-scientists. document.
About Issues in Ecology Cover photo credits, clockwise from top left:
Issues in Ecology is designed to report, in lan- Nadine Cavender, Nadine Cavender, Nadine Cavender,
guage understandable by non-scientists, the consensus Claude Cavender, Jr., Nadine Cavender, unknown, Claude
of a panel of scientific experts on issues relevant to the Cavender, Jr., and unknown.
16