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9

Ecosysrem Services: benefirs


Supplied ro Human Socieries
by Narural Ecosysrems
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Issoes |n Lco|ogy Nomher 2 Spr|ng 1997


1
Lcosystem Serv|ces:
8enet|ts Sopp||eJ to Homan
Soc|et|es hy Natora| Lcosystems
SUMMAkY
Human socieries derive many essenrial goods from narural ecosysrems, including seafood, game animals, fodder,
fuelwood, rimber, and pharmaceurical producrs. These goods represenr imporranr and familiar parrs of rhe economy.
vhar has been less appreciared unril recenrly is rhar narural ecosysrems also perform fundamenral lifesupporr services
wirhour which human civilizarions would cease ro rhrive. These include rhe purificarion of air and warer, deroxificarion
and decomposirion of wasres, regularion of climare, regenerarion of soil ferriliry, and producrion and mainrenance of
biodiversiry, from which key ingredienrs of our agriculrural, pharmaceurical, and indusrrial enrerprises are derived. This
array of services is generared by a complex inrerplay of narural cycles powered by solar energy and operaring across a
wide range of space and rime scales. The process of wasre disposal, for example, involves rhe life cycles of bacreria as well
as rhe planerwide cycles of major chemical elemenrs such as carbon and nirrogen. Such processes are worrh many
rrillions of dollars annually. Yer because mosr of rhese benefirs are nor rraded in economic markers, rhey carry no price
rags rhar could alerr sociery ro changes in rheir supply or dereriorarion of underlying ecological sysrems rhar generare
rhem. because rhrears ro rhese sysrems are increasing, rhere is a crirical need for idenrificarion and moniroring of
ecosysrem services borh locally and globally, and for rhe incorporarion of rheir value inro decisionmaking processes.
Hisrorically, rhe narure and value of Earrh`s life supporr sysrems have largely been ignored unril rheir disruprion or
loss highlighred rheir imporrance. |or example, deforesrarion has belaredly revealed rhe crirical role foresrs serve in
regularing rhe warer cycle in parricular, in mirigaring floods, droughrs, rhe erosive forces of wind and rain, and silring
of dams and irrigarion canals. Today, escalaring impacrs of human acriviries on foresrs, werlands, and orher narural
ecosysrems imperil rhe delivery of such services. The primary rhrears are land use changes rhar cause losses in biodiversiry
as well as disruprion of carbon, nirrogen, and orher biogeochemical cycles: humancaused invasions of exoric species:
releases of roxic subsrances: possible rapid climare change: and deplerion of srrarospheric ozone.
based on available scienrific evidence, we are cerrain rhar:
Ecosysrem services are essenrial ro civilizarion.
Ecosysrem services operare on such a grand scale and in such inrricare and lirrleexplored ways rhar mosr could nor
be replaced by rechnology.
Human acriviries are already impairing rhe flow of ecosysrem services on a large scale.
|f currenr rrends conrinue, humaniry will dramarically alrer virrually all of Earrh`s remaining narural ecosysrems wirhin
a few decades.
|n addirion, based on currenr scienrific evidence, we are confidenr rhar:
Many of rhe human acriviries rhar modify or desrroy narural ecosysrems may cause dereriorarion of ecological
services whose value, in rhe long rerm, dwarfs rhe shorrrerm economic benefirs sociery gains from rhose acriviries.
Considered globally, very large numbers of species and popularions are required ro susrain ecosysrem services.
The funcrioning of many ecosysrems could be resrored if appropriare acrions were raken in rime.
ve believe rhar land use and developmenr policies should srrive ro achieve a balance berween susraining viral
ecosysrem services and pursuing rhe worrhy shorrrerm goals of economic developmenr.
1J
Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
Lcosystem Serv|ces: 8enet|ts Sopp||eJ
to Homan Soc|et|es hy Natora| Lcosystems
by
Crerchen C. Daily, Susan Alexander, Paul R. Ehrlich,
Larry Coulder, 1ane Lubchenco, Pamela A. Marson, Harold A. Mooney,
Sandra Posrel, Srephen H. Schneider, David Tilman, Ceorge M. voodwell
INTkODUCTION
Many socieries roday have rechnological capa
biliries undreamed of in cenruries pasr. Their cirizens
have such a global command of resources rhar even foods
flown in fresh from all over rhe planer are raken for
granred, and daily menus are decoupled from rhe limira
rions of regional growing seasons and soils. These de
velopmenrs have focused so much arrenrion upon
humanengineered and exoric sources of fulfillmenr rhar
rhey diverr arrenrion from rhe local biological underpin
nings rhar remain essenrial ro economic prosperiry and
orher aspecrs of our wellbeing.
These biological underpinnings are encompassed
in rhe phrase ecosysrem services, which refers ro a wide
range of condirions and processes rhrough which naru
ral ecosysrems, and rhe species rhar are parr of rhem,
help susrain and fulfill human life. These services main
rain biodiversiry and rhe producrion of ecosysrem goods,
such as seafood, wild game, forage, rimber, biomass fu
els, narural fibers, and many pharmaceuricals, indusrrial
producrs, and rheir precursors. The harvesr and rrade of
rhese goods represenr imporranr and familiar parrs of
rhe human economy. |n addirion ro rhe producrion of
goods, ecosysrem services supporr life rhrough (Holdren
and Ehrlich 194: Ehrlich and Ehrlich 19S1j:
purificarion of air and warer.
mirigarion of droughrs and floods.
generarion and preservarion of soils and renewal of
rheir ferriliry.
deroxificarion and decomposirion of wasres.
pollinarion of crops and narural vegerarion.
dispersal of seeds.
cycling and movemenr of nurrienrs.
conrrol of rhe vasr majoriry of porenrial agriculrural
pesrs.
mainrenance of biodiversiry.
prorecrion of coasral shores from erosion by waves.
prorecrion from rhe sun`s harmful ulrravioler rays.
parrial srabilizarion of climare.
moderarion of wearher exrremes and rheir impacrs.
provision of aesrheric beaury and inrellecrual srimu
larion rhar lifr rhe human spirir.
Alrhough rhe disrincrion berween narural" and
humandominared" ecosysrems is becoming increasingly
blurred, we emphasize rhe narural end of rhe specrrum,
for rhree relared reasons. |irsr, rhe services flowing from
narural ecosysrems are grearly undervalued by sociery.
|or rhe mosr parr, rhey are nor rraded in formal markers
and so do nor send price signals rhar warn of changes in
rheir supply or condirion. |urrhermore, few people are
conscious of rhe role narural ecosysrem services play in
||gore 1Aspen (Pcpa|as trema|c|desj
foresr in Colorado, filrering and pu
rifying air and warer.
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Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
||gore 2voman carrying rreerrunk for boar
making in a fishing village on Chiloe |sland, Chile.
Narural foresrs remain an imporranr source of
wood for consrrucrion, fuel, and orher uses.
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generaring rhose ecosysrem goods rhar are rraded in rhe
markerplace. As a resulr, rhis lack of awareness helps
dri ve rhe conversi on of narural ecosysrems ro
humandominared sysrems (e.g., whearlands or oil palm
fieldsj, whose economic value can be expressed, ar leasr
in parr, in srandard currency. The second reason ro focus
on narural ecosysrems is rhar many humaniniriared dis
ruprions of rhese sysrems such as inrroducrions of ex
oric species, exrincrions of narive species, and alrerarion
of rhe gaseous composirion of rhe armosphere rhrough
fossil fuel burning are difficulr or impossible ro reverse
on any rime scale relevanr ro sociery. Third, if awareness
is nor increased and currenr rrends conrinue, humaniry
will dramarically alrer Earrh`s remaining narural ecosys
rems wirhin a few decades (Daily 199a, bj.
The lack of arrenrion ro rhe viral role of narural
ecosysrem services is easy ro undersrand. Humaniry came
inro being afrer mosr ecosysrem services had been in
operarion for hundreds of millions ro billions of years.
These services are so fundamenral ro life rhar rhey are
easy ro rake for granred, and so large in scale rhar ir is
hard ro imagine rhar human acriviries could irreparably
disrupr rhem. Perhaps a rhoughr experimenr rhar re
moves rhese services from rhe familiar backdrop of rhe
Earrh is rhe besr way ro illusrrare borh rhe imporrance
and complexiry of ecosysrem services, as well as how
illequipped humans are ro recreare rhem. |magine, for
example, human beings rrying ro colonize rhe moon.
Assume for rhe sake of argumenr rhar rhe moon had
already miraculously acquired some of rhe basic condi
rions for supporring human life, such as an armosphere,
a climare, and a physical soil srrucrure similar ro rhose
on Earrh. The big quesrion facing human colonisrs would
rhen be, which of Earrh`s millions of species would need
ro be rransporred ro rhe moon ro make rhar srerile sur
face habirable'
One could rackle rhar quesrion sysremarically
by firsr choosing from among all rhe species exploired
direcrly for food, drink, spices, fiber, rimber, pharmaceu
ricals, and indusrrial producrs such as waxes, rubber, and
oils. Even if one were highly selecrive, rhe lisr could amounr
ro hundreds or even rhousands of species. And rhar would
only be a srarr, since one would rhen need ro consider
which species are crucial ro supporring rhose used di
recrly: rhe bacreria, fungi, and inverrebrares rhar help
make soil ferrile and break down wasres and organic
marrer: rhe insecrs, bars, and birds rhar pollinare flow
ers: and rhe grasses, herbs, and rrees rhar hold soil in
place, regulare rhe warer cycle, and supply food for ani
mals. The clear message of rhis exercise is rhar no one
knows which combinarions of species or even approxi
marely how many are required ro susrain human life.
Rarher rhan selecring species direcrly, one mighr
rry anorher approach: Lisring rhe ecosysrem services
needed by a lunar colony and rhen guessing ar rhe rypes
and numbers of species required ro perform each. Yer
derermining which species are crirical ro rhe funcrioning
of a parricular ecosysrem service is no simple rask. Ler
us rake soil ferriliry as an example. Soil organisms are
crucial ro rhe chemical conversion and physical rransfer
of essenrial nurrienrs ro higher planrs. bur rhe abun
dance of soil organisms is absolurely sraggering. Under
a squareyard of pasrure in Denmark, for insrance, rhe
soil is inhabired by roughly 50,000 small earrhworms
and rheir relarives, 50,000 insecrs and mires, and nearly
1J million roundworms. And rhar rally is only rhe begin
ning. The number of soil animals is riny compared ro rhe
number of soil microorganisms: a pinch of ferrile soil may
conrain over J0,000 prorozoa, 50,000 algae, 400,000
fungi , and bi l l i ons of i ndi vi dual bacreri a
(OvergaardNielsen 1955: Rouarr and Karznelson 1961:
Chanway 199Jj. vhich musr colonisrs bring ro rhe moon
ro assure lush and conrinuing planr growrh, soil renewal,
wasre disposal, and so on' Mosr of rhese soildwelling
species have never been subjecred ro even cursory in
specrion: no human eye has ever blinked ar rhem rhrough
4
Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
||gore 3Alpaca grazing on rhe Chilean alri
plano. Crassland ecosysrems are an impor
ranr source of animal producrs: rhey are also
rhe original habirar of mosr domesric animals
and many crops, such as whear, barley, and
oars.
a microscope, no human hand has ever ryped our a name
or descriprion of rhem, and mosr human minds have never
spenr a momenr reflecring on rhem. Yer rhe sobering
facr is, as E. O. vilson pur ir: rhey don`r need us, bur we
need rhem (vilson 19Sj.
THL CHAkACTLk O| LCOSYSTLM SLkvICLS
Moving our arrenrion from rhe moon back ro Earrh, ler
us look more closely ar rhe services narure performs on
rhe only planer we know rhar is habirable. Ecosysrem
services and rhe sysrems rhar supply rhem are so inrer
connecred rhar any classificarion of rhem is necessarily
rarher arbirrary. Here we briefly explore a suire of
overarching services rhar operare in ecosysrems world
wide.
Prcdact|ca c[ Fccsqstcm Cccds
Humaniry obrains from narural ecosysrems an
array of ecosysrem goods organisms and rheir parrs
and producrs rhar grow in rhe wild and rhar are used
direcrly for human benefir. Many of rhese, such as fishes
and animal producrs, are commonly rraded in economic
markers. The annual world fish carch, for example,
amounrs ro abour 100 million merric rons and is valued
ar berween 550 billion and 5100 billion: ir is rhe leading
source of animal prorein, wirh over J0% of rhe popula
rion in Africa and Asia dependenr on fish as rheir pri
mary source of prorein (UN|AO 199Jj. The commercial
harvesr of freshwarer fish worldwide in 1990 roraled
approximarely 14 million rons and was valued ar abour
5S.J billion (UN|AO 1994j. |nreresringly, rhe value of
rhe freshwarer sporr fishery in rhe U.S. alone grearly
exceeds rhar of rhe global commercial harvesr, wirh di
recr expendirures in 1991 roraling abour 516 billion.
vhen rhis is added ro rhe value of rhe employmenr gen
erared by sporr fishing acriviries, ir raises rhe roral ro
546 billion (|elder and Nickum 199J, cired in Posrel and
Carpenrer 199j. The furure of rhese fisheries is in ques
rion, however, because fish harvesrs have approached or
exceeded susrainable levels virrually everywhere. Nine
of rhe world`s major marine fishing areas are in decline
due ro overfishing, pollurion, and habirar desrrucrion.
(UN|AO 199J: Kaufman and Dayron 199j.
Turning our arrenrion ro rhe land, grasslands are
an imporranr source of markerable goods, including ani
mals used for labor (horses, mules, asses, camels, bul
locks, erc.j and rhose whose parrs or producrs are con
sumed (as mear, milk, wool, and learherj. Crasslands
were also imporranr as rhe original source habirar for
mosr domesric animals such as carrle, goars, sheep, and
horses, as well as many crops, such as whear, barley,
rye, oars, and orher grasses (Sala and Paruelo 199j. |n
a wide variery of rerresrrial habirars, people hunr game
animals such as warerfowl, deer, moose, elk, fox, boar
and orher wild pigs, rabbirs, and even snakes and mon
keys. |n many counrries, game mear forms an imporranr
parr of local diers and, in many places, hunring is an
economically and culrurally imporranr sporr.
Narural ecosysrems also produce vegerarion used direcrly
by humans as food, rimber, fuelwood, fiber, pharmaceu
ricals and indusrrial producrs. |ruirs, nurs, mushrooms,
honey, orher foods, and spices are exrracred from many
foresr species. vood and orher planr marerials are used
in rhe consrrucrion of homes and orher buildings, as well
as for rhe manufacrure of furnirure, farming implemenrs,
paper, clorh, rharching, rope, and so on. Abour 15 per
cenr of rhe world`s energy consumprion is supplied by
fuelwood and orher planr marerial: in developing coun
rries, such biomass" supplies nearly 40 percenr of en
ergy consumprion (Hall er al. 199Jj, alrhough rhe por
ri on of rhi s deri ved from narural rarher rhan
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As described in rhe previous secrion, biodiversiry
is a direcr source of ecosysrem goods. |r also supplies
rhe generic and biochemical resources rhar underpin our
currenr agriculrural and pharmaceurical enrerprises and
may allow us ro adapr rhese viral enrerprises ro global
change. Our abiliry ro increase crop producriviry in rhe
face of new pesrs, diseases, and orher srresses has de
pended heavily upon rhe rransfer ro our crops of genes
from wild crop relarives rhar confer resisrance ro rhese
challenges. Such exrracrions from biodiversiry`s generic
library" accounr for annual increases in crop producriv
iry of abour 1 percenr, currenrly valued ar 51 billion (NRC
199Jj. biorechnology now makes possible even grearer
use of rhis narural srorehouse of ge
neric diversiry via rhe rransfer ro crops
of genes from any kind of organism
nor simply crop relarivesand ir
promises ro play a major role in fu
rure yield increases. by rhe rurn of
rhe cenrury, farmlevel sales of rhe
producrs of agriculrural biorechnol
ogy, jusr now enrering rhe marker
place, are expecred ro reach ar leasr
510 billion per year (vorld bank
1991, cired in Reid er al. 1996j.
|n addirion ro susraining rhe produc
rion of convenrional crops, rhe
biodiversiry in narural ecosysrems may
include many porenrial new foods.
Human beings have urilized around
,000 planr species for food over rhe
course of hisrory and anorher 0,000
planrs are known ro have edible parrs
(vilson 19S9j. Only abour 150 food
planrs have ever been culrivared on a
large scale, however. Currenrly, SJ
planr species conrribure 90 percenr
of narional percapira supplies of food
pl anrs (PrescorrAl l en and
PrescorrAllen 1990j, alrhough a much smaller number
of rhese supply rhe bulk of rhe calories humans consume.
Many orher species, however, appear more nurririous or
berrer suired ro rhe growing condirions rhar prevail in
imporranr regions rhan rhe srandard crops rhar domi
nare world food supply roday. because of increasing
salinizarion of irrigared croplands and rhe porenrial for
rapid climare change, for insrance, furure food securiry
may come ro depend on droughr and salrroleranr vari
eries rhar now play compararively minor roles in agricul
rure.
5
Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
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||gore 4Harpoonwhaling in |lores, |n
donesia. The oceans are a key source
of animal prorein for rhe human popu
larion.
humandominared ecosysrems is undocumenred. |n addi
rion, narural producrs exrracred from many hundreds of
species conrribure diverse inpurs ro indusrry: gums and
exudares, essenrial oils and flavorings, resins and oleo
resins, dyes, rannins, vegerable fars and waxes, insecri
cides, and mulrirudes of orher compounds (Myers 19SJ:
Leung and |osrer 1996j. The availabiliry of mosr of
rhese narural producrs is in decline due ro ongoing habi
rar conversion.
Ccacrat|ca aad Ma|atcaaacc c[ 8|cd|ucrs|tq
biological diversiry, or biodiversiry for shorr, re
fers ro rhe variery of life forms ar all levels of organiza
rion, from rhe molecular ro rhe land
scape level. biodiversiry is generared
and mainrained in narural ecosysrems,
where organisms encounrer a wide
variery of living condirions and chance
evenrs rhar shape rheir evolurion in
unique ways. Our of convenience or
necessiry, biodiversiry is usually quan
rified in rerms of numbers of species,
and rhis perspecrive has grearly influ
enced conservarion goals. |r is im
porranr ro remember, however, rhar
rhe benefirs rhar biodiversiry supplies
ro humaniry are delivered rhrough
popularions of species residing in liv
ing communiries wirhin specific physi
cal serrings in orher words, rhrough
complex ecological sysrems, or eco
sysrems (Daily and Ehrlich 1995j. |or
human beings ro realize mosr of rhe
aesrheric, spirirual, and economic ben
efirs of biodiversiry, narural ecosys
rems musr rherefore be accessible. The
conrinued exisrence of coniferous rree
species somewhere in rhe world would
nor help rhe inhabiranrs of a rown in
undared by flooding because of rhe clearing of a pine
foresr upsrream. Cenerally, rhe flow of ecosysrem goods
and services in a region is derermined by rhe rype, spa
rial layour, exrenr, and proximiry of rhe ecosysrems sup
plying rhem. because of rhis, rhe preservarion of only
one minimum viable popularion of each nonhuman spe
cies on Earrh in zoos, boranical gardens, and rhe world`s
legally prorecred areas would nor susrain life as we know
ir. |ndeed, such a srraregy, raken ro exrreme, would lead
ro collapse of rhe biosphere, along wirh irs life supporr
services.
6
Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
||gore 5Trapping and releasing burrerflies
in a mixedagriculrure landscape in Cosra
Rica. Moniroring rhe impacr of human
acriviries on biodiversiry and ecosysrem
services is needed worldwide: burrerflies
may be useful indicarors for moniroring.
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Turning ro medicinal resources, a recenr survey
showed rhar of rhe rop 150 prescriprion drugs used in
rhe Unired Srares, 11S are based on narural sources:
4% on planrs, 1S% on fungi, 5% on bacreria, and J%
on one verrebrare (snakej species. Nine of rhe rop ren
drugs in rhis lisr are based on narural planr producrs
(Crifo and Rosenrhal, in press, as cired in Dobson 1995j.
The commercial value of pharmaceuricals in rhe devel
oped narions exceeds 540 billion per year (Principe 19S9j.
Looking ar rhe global picrure, approximarely S0% of rhe
human popularion relies on rradirional medical sysrems,
and abour S5% of rradirional medicine involves rhe use
of planr exrracrs (|arnsworrh er al. 19S5j.
Saving only a single popula
rion of each species could have an
orher cosr. Differenr popularions of
rhe same species may produce differ
enr rypes or quanriries of defensive
chemicals rhar have porenrial use as
pharmaceuri cal s or pesri ci des
(McCormick er al. 199Jj: and rhey
may exhibir differenr rolerances ro
envi ronmenral srresses such as
droughr or soil saliniry. |or example,
rhe developmenr of penicillin as a
rherapeuric anribioric rook a full 15
years afrer Alexander |leming`s fa
mous discovery of ir in common bread
mold. |n parr, rhis was because sci
enrisrs had grear difficulry producing,
exrracring, and purifying rhe sub
srance in needed quanriries. One key
ro obraining such quanriries was rhe
discovery, afrer a worldwide search,
of a popularion of |leming`s mold rhar
produced more penicillin rhan rhe
original (Dowling 19j. Similarly,
planr popularions vary in rheir abiliry
ro resisr pesrs and disease, rrairs im
porranr in agriculrure. Many rhousands of varieries of
rice from differenr locarions were screened ro find one
wirh resisrance ro grassy srunr virus, a disease rhar posed
a serious rhrear ro rhe world`s rice crop (Myers 19SJj.
Despire numerous examples like rhese, many of rhe lo
caliries rhar harbor wild relarives of crops remain unpro
recred and heavily rhrearened.
C||matc aad l|[c
Earrh`s climare has flucruared rremendously since
humaniry came inro being. Ar rhe peak of rhe lasr ice
age J0,000 years ago, for example, much of Europe and
Norrh America were covered by milerhick ice sheers.
vhile rhe global climare has been relarively srable since
rhe invenrion of agriculrure around 10,000 years ago,
periodic shifrs in climare have affecred human acriviries
and serrlemenr parrerns. Even relarively recenrly, from
15501S50, Europe was significanrly cooler during a
period known as rhe Lirrle |ce Age. Many of rhese changes
in climare are rhoughr ro be caused by alrerarions in
Earrh`s orbiral rorarion or in rhe energy ourpur of rhe
sun, or even by evenrs on rhe Earrh irselfsudden per
rurbarions such as violenr volcanic eruprions and asrer
oid impacrs or more gradual recronic evenrs such as rhe
uplifr of rhe Himalayas. Remarkably,
climare has been buffered enough
rhrough all rhese changes ro susrain
life for ar leasr J.5 billion years
(Schneider and Londer 19S4j. And
life irself has played a role in rhis buff
ering.
Climare, of course, plays a
major role in rhe evolurion and disrri
burion of life over rhe planer. Yer mosr
scienrisrs would agree rhar life irself
is a principal facror in rhe regularion
of global climare, helping ro offser rhe
effecrs of episodic climare oscillarions
by responding in ways rhar alrer rhe
greenhouse gas concenrrarions in rhe
armosphere. |or insrance, narural eco
sysrems may have helped ro srabilize
climare and prevenr overhearing of rhe
Earrh by removing more of rhe green
house gas carbon dioxide from rhe
armosphere as rhe sun grew brighrer
over millions of years (Alexander er
al. 199j. Life may also exerr a de
srabilizing or posirive feedback rhar
reinforces climare change, parricularly
during rransirions berween inrerglacial periods and ice
ages. One example: vhen climaric cooling leads ro drops
in sea level, conrinenral shelves are exposed ro wind and
rain, causing grearer nurrienr runoff ro rhe oceans. These
nurrienrs may ferrilize rhe growrh of phyroplankron, many
of which form calcium carbonare shells. |ncreasing rheir
popularions would remove more carbon dioxide from rhe
oceans and rhe armosphere, a mechanism rhar should
furrher cool rhe planer. Living rhings may also enhance
warming rrends rhrough such acriviries as speeding up
microbial decomposirion of dead organic marrer, rhus
7
Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
||gore 6bark of rhe Pacific yew rree (Taxas hrec|jc||aj,
which is rhe source of rhe new anricancer drug, raxol,
villamerre Narional |oresr, Oregon.
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||gore 7Herbal pharmacisr in Dali, Yunnan Province,
China. An esrimared S0 percenr of rhe world`s popu
larion relies on narural medicinal producrs.
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releasing carbon dioxide ro rhe armosphere (Schneider
and bosron 1991: Allegre and Schneider 1994j. The
relarive influence of life`s srabilizing and desrabilizing feed
backs remains uncerrain: whar is clear is rhar climare
and narural ecosysrems are righrly coupled, and rhe sra
bi l i ry of rhar coupl ed sysrem i s an i mporranr
ecosysremservice.
besides rheir impacr on rhe armosphere, ecosys
rems also exerr direcr physical influences rhar help ro
moderare regional and local wearher. |or insrance, rran
spirarion (release of warer vapor from rhe leavesj of planrs
in rhe morning causes rhundersrorms in rhe afrernoon,
limiring borh moisrure loss from rhe region and rhe rise
in surface remperarure. |n rhe Amazon, for example,
50% of rhe mean annual rainfall is recycled by rhe foresr
irself via evaporranspirarionrhar is, evaporarion from
wer leaves and soil combined wirh rranspirarion (Salari
19Sj. Amazon deforesrarion could so dramarically re
duce roral precipirarion rhar rhe foresr mighr be unable
ro reesrablish irself following complere desrrucrion (Shukla
er al. 1990j. Temperarure exrremes are also moderared
by foresrs, which provide shade and surface cooling and
also acr as insularors, blocking searing winds and rrap
ping warmrh by acring as a local greenhouse agenr.
M|t|qat|ca c[ F|ccds aad Drcaqhts
An enormous amounr of warer, abour 119,000
cubic kilomerers, is rained annually onro rhe Earrh`s land
surfaceenough ro cover rhe land ro an average deprh
of 1 merer (Shiklomanov 199Jj. Much of rhis warer is
soaked up by soils and gradually mered our ro planr roors
or inro aquifers and surface srreams. Thus, rhe soil irself
slows rhe rush of warer off rhe land in flash floods. Yer
bare soil is vulnerable. Planrs and planr lirrer shield rhe
soil from rhe full, desrrucrive force of raindrops and hold
ir in place. vhen landscapes are denuded, rain com
pacrs rhe surface and rapidly rurns soil ro mud (espe
cially if ir has been loosened by rillagej: mud clogs sur
face caviries in rhe soil, reduces infilrrarion of warer, in
creases runoff, and furrher enhances clogging. Derached
soil parricles are splashed downslope and carried off by
running warer (Hillel 1991j.
Erosion causes cosrs nor only ar rhe sire where
soil is losr bur also in aquaric sysrems, narural and
humanmade, where rhe marerial accumulares. Local
cosrs of erosion include losses of producrion porenrial,
diminished infilrrarion and warer availabiliry, and losses
of nurrienrs. Downsrream cosrs may include disrupred
or lower qualiry warer supplies: silrarion rhar impairs drain
age and mainrenance of navigable river channels, har
bors, and irrigarion sysrems: increased frequency and
severiry of floods: and decreased porenrial for hydroelec
rric power as reservoirs fill wirh silr (Pimenrel er al. 1995j.
vorldwide, rhe replacemenr cosr of reservoir capaciry
losr ro silrarion is esrimared ar 56 billion per year.
|n addirion ro prorecring soil from erosion, living
vegerarionwirh irs deep roors and aboveground evapo
raring surfacealso serves as a gianr pump, rerurning
warer from rhe ground inro rhe armosphere. Clearing of
planr cover disruprs rhis link in rhe warer cycle and leads
Scru|ccs Sapp||cd bq Sc||
Soil represenrs an imporranr componenr of a
narion`s assers, one rhar rakes hundreds ro hundreds of
rhousands of years ro build up and yer very few years ro
be losr. Some civilizarions have drawn grear srrengrh
from ferrile soil: conversely, rhe loss of producriviry
rhrough mismanagemenr is rhoughr ro have ushered many
once flourishing socieries ro rheir ruin (Adams 19S1j.
Today, soil degradarion induced by human acriviries af
flicrs nearly J0 percenr of rhe Earrh`s vegerared land
surface (Oldeman er al. 1990j.
|n addirion ro moderaring rhe warer cycle, as described
above, soil provides five orher inrerrelared services (Daily
er al. 199j. |irsr, soil shelrers seeds and provides physi
cal supporr as rhey sprour and marure inro adulr planrs.
The cosr of packaging and sroring seeds and of anchor
ing planr roors would be enormous wirhour soil.
Humanengineered hydroponic sysrems can grow planrs
in rhe absence of soil, and rheir cosr provides a lower
bound ro help assess rhe value of rhis service. The cosrs
of physical supporr rrays and srands used in such opera
rions roral abour US555,000 per hecrare (for rhe Nurri
enr |ilm Technique Sysrems: |AO 1990j.
Second, soil rerains and delivers nurrienrs ro
planrs. Tiny soil parricles (less rhan J microns in diam
ererj, which are primarily birs of humus and clays, carry
a surface elecrrical charge rhar is generally negarive. This
properry holds posirively charged nurrienrscarions such
as calcium and magnesiumnear rhe surface, in prox
imiry ro planr roors, allowing rhem ro be raken up gradu
ally. Orherwise, rhese nurrienrs would quickly be leached
away. Soil also acrs as a buffer in rhe applicarion of
ferrilizers, holding onro rhe ferrilizer ions unril rhey are
8
Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
||gore 8Early summer in
rhe Colorado Rockies.
These subalpine foresrs
mirigare flood, droughr,
and remperarure exrremes:
rhey soak up rain and
snowmelr and mere ir our
gradually ro srreams and
ro rhe armosphere, crear
i ng cool i ng afrernoon
rhundersrorms.
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ro porenrially large increases in surface runoff, along wirh
nurrienr and soil loss. A classic example comes from rhe
experimenral clearing of a New Hampshire foresr, where
herbicide was applied ro prevenr regrowrh for a Jyear
period afrer rhe clearing. The resulr was a 40 percenr
increase in average srream flow. During one fourmonrh
period of rhe experimenr, runoff was more rhan 5 rimes
grearer rhan before rhe clearing (bormann 196Sj. On a
much larger scale, exrensive deforesrarion in rhe Hima
layan highlands appears ro have exacerbared recenr flood
ing in bangladesh, alrhough rhe relarive roles of human
and narural forces remain debarable (|ves and Messerli
19S9j. |n addirion, some regions of rhe world, such as
parrs of Africa, are experiencing an increased frequency
and severiry of droughr, possibly associared wirh exren
sive deforesrarion.
verlands are parricularly wellknown for rheir role
in flood conrrol and can ofren reduce rhe need ro con
srrucr flood conrrol srrucrures. |loodplain foresrs and
high salr marshes, for example, slow rhe flow of floodwa
rers and allow sedimenrs ro be deposired wirhin rhe flood
plain rarher rhan washed inro downsrream bays or oceans.
|n addirion, isolared werlands such as prairie porholes in
rhe Midwesr and cypress ponds in rhe Sourheasr, serve
as derenrion areas during rimes of high rainfall, delaying
sarurarion of upland soils and overland flows inro rivers
and rhereby damping peak flows. Reraining rhe inregriry
of rhese werlands by leaving vegerarion, soils, and naru
ral warer regimes inracr can reduce rhe severiry and du
rarion of flooding along rivers (Ewel 199j. A relarively
small area of rerained werland, for example, could have
largely prevenred rhe severe flooding along rhe Missis
sippi River in 199J.
9
Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
||gore 9bacreria (Bradqrh|zch|am japca|camj
in a soybean roor nodule cell, magnified J,550
rimes. These bacreria fix armospheric nirro
gen inro a form rhar can be urilized by planrs.
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required by planrs. Hydroponic sysrems supply warer
and nurrienrs ro planrs wirhour need of soil, bur rhe mar
gin for error is much smallereven small excesses of
nurrienrs applied hydroponically can be lerhal ro planrs.
|ndeed, ir is a complex underraking ro regulare rhe nurri
enr concenrrarions, pH, and saliniry of rhe nurrienr solu
rion in hydroponic sysrems, as well as rhe air and solu
rion remperarure, humidiry, lighr, pesrs, and planr dis
eases. vorldwide, rhe area under hydroponic culrure is
only a few rhousand hecrares and is unlikely ro grow
significanrly in rhe foreseeable furure: by conrrasr, glo
bal cropped area is abour 1.4 billion hecrares (USDA
199Jj.
Third, soil plays a cenrral
role in rhe decomposirion of dead
organic marrer and wasres, and rhis
decomposirion process also renders
harmless many porenrial human
parhogens. People generare a rre
mendous amounr of wasre, includ
ing household garbage, indusrrial
wasre, crop and foresrry residues,
and sewage from rheir own popula
rions and rheir billions of domesri
cared animals. A rough approxima
rion of rhe amounr of dead organic
marrer and wasre (mosrly agricul
rural residuesj processed each year
is 1J0 billion merric rons, abour J0
percenr of which is associared wirh
human acriviries (derived from
virousek er al. 19S6j. |orrunarely,
rhere is a wide array of decompos
ing organismsranging from vul
rures ro riny bacreriarhar exrracr
energy from rhe large, complex organic molecules found
in many rypes of wasre. Like assemblyline workers, di
verse microbial species process rhe parricular compounds
whose chemical bonds rhey can cleave and pass along ro
orher species rhe end producrs of rheir specialized reac
rions. Many indusrrial wasres, including soaps, derergenrs,
pesricides, oil, acids, and paper, are deroxified and de
composed by organisms in narural ecosysrems if rhe con
cenrrarion of wasre does nor exceed rhe sysrem`s capac
iry ro rransform ir. Some modern wasres, however, are
virrually indesrrucrible, such as some plasrics and rhe
breakdown producrs of rhe pesricide DDT.
The simple inorganic chemicals rhar resulr from
narural decomposirion are evenrually rerurned ro planrs
as nurrienrs. Thus, rhe decomposirion of wasres and rhe
recycling of nurrienrsrhe fourrh service soils provide
are rwo aspecrs of rhe same process. The ferriliry of
soilsrhar is, rheir abiliry ro supply nurrienrs ro planrs
is largely rhe resulr of rhe acriviries of diverse species of
bacreria, fungi, algae, crusracea, mires, rermires, spring
rails, millipedes, and worms, all of which, as groups, play
imporranr roles. Some bacreria are responsible for fix
ing" nirrogen, a key elemenr in proreins, by drawing ir
our of rhe armosphere and converring ir ro forms usable
by planrs and, ulrimarely, human beings and orher ani
mals. Cerrain rypes of fungi play exrremely imporranr
roles in supplying nurrienrs ro many kinds of rrees. Earrh
worms and anrs acr as mechani
cal blenders," breaking up and mix
ing planr and microbial marerial and
orher marrer (1enny 19S0j. |or
example, as much as 10 merric
ronnes of mareri al may pass
rhrough rhe bodies of earrhworms
on a hecrare of land each year, re
sulring in nurrienr rich casrs" rhar
enhance soil srabiliry, aerarion, and
drainage (Lee 19S5j.
|inally, soils are a key fac
ror in regularing rhe Earrh`s major
elemenr cyclesrhose of carbon,
nirrogen, and sulfur. The amounr
of carbon and nirrogen srored in
soils dwarfs rhar in vegerarion, for
example. Carbon in soils is nearly
double (1.S rimesj rhar in planr
marrer, and nirrogen in soils is abour
1S rimes grearer (Schlesinger
1991j. Alrerarions in rhe carbon
and nirrogen cycles may be cosrly
over rhe long rerm, and in many cases, irreversible on a
rime scale of inreresr ro sociery. |ncreased fluxes of car
bon ro rhe armosphere, such as occur when land is con
verred ro agriculrure or when werlands are drained, con
rribure ro rhe buildup of key greenhouse gases, namely
carbon dioxide and merhane, in rhe armosphere
(Schlesinger 1991j. Changes in nirrogen fluxes caused
by producrion and use of ferrilizer, burning of wood and
orher biomass fuels, and clearing of rropical land lead ro
increasing armospheric concenrrarions of nirrous oxide,
anorher porenr greenhouse gas rhar is also involved in
rhe desrrucrion of rhe srrarospheric ozone shield. These
and orher changes in rhe nirrogen cycle also resulr in
acid rain and excess nurrienr inpurs ro freshwarer sys
rems, esruaries, and coasral marine warers. This nurrienr
10
||gore 10Sonoran bumble bee (Bcmhas scacrasj pollinar
ing a flower.
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Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
influx causes eurrophicarion of aquaric ecosysrems and
conraminarion of drinking warer sourcesborh surface
and ground warer by high levels of nirrarenirrogen
(virousek er al. 199j.
Pc|||aat|ca
Animal pollinarion is required for rhe successful
reproducrion of mosr flowering planrs. Abour JJ0,000
our of an esrimared J40,000 species of planrs for which
rhe mode of pollinarion has been recorded require an
animal such as a bee or hummingbird ro accomplish rhis
viral rask. This includes borh wild planrs and abour 0
percenr of rhe agriculrural crop species rhar feed rhe
world. Over 100,000 differenr animal speciesinclud
ing bars, bees, beerles, birds, burrerflies, and fliesare
known ro provide rhese free pollinarion services rhar as
sure rhe perperuarion of planrs in our croplands, back
yard gardens, rangelands,
meadows and foresrs. |n
rurn, rhe conrinued availabil
iry of rhese pollinarors de
pends on rhe exisrence of a
wide variery of habirar rypes
needed for rheir feeding, suc
cessful breedi ng, and
complerion of rheir life cycles
(Nabhan and buchmann
199j.
One rhird of human
food is derived from planrs
pollinared by wild pollinarors.
virhour narural pollinarion
services, yields of imporranr crops would decline precipi
rously and many wild planr species would become ex
rincr. |n rhe Unired Srares alone, rhe agriculrural value
of wild, narive pollinarorsrhose susrained by narural
habirars adjacenr ro farmlands is esrimared in rhe bil
lions of dollars per year. Pollinarion by honey bees, origi
nally imporred from Europe, is exrremely imporranr as
well, bur rhese bees are presenrly in decline, enhancing
rhe imporrance of pollinarors from narural ecosysrems.
Managemenr of rhe honey bee in rhe New vorld is cur
renrly rhrearened by rhe movemenr of, and hybridizarion
wirh, an aggressive African srrain of honey bee rhar was
accidenrally released in brazil in 1956. Diseases of honey
bee colonies are also causing a marked decline in rhe
number of managed colonies. Meanwhile, rhe diversiry
of narural pollinarors available ro borh wild and domesri
cared planrs is diminishing: more rhan 60 genera of pol
linarors include species now considered ro be rhrearened,
endangered or exrincr (buchmann and Nabhan 1996j.
Natara| Pcst Ccatrc| Scru|ccs
Humaniry`s comperirors for food, rimber, corron,
and orher fibers are called pesrs, and rhey include nu
merous herbivorous insecrs, rodenrs, fungi, snails, nema
rodes, and viruses. These pesrs desrroy an esrimared J5
ro 50 percenr of rhe world`s crops, eirher before or afrer
harvesr (Pimenrel er al. 19S9j. |n addirion, numerous
weeds compere direcrly wirh crops for warer, lighr, and
soil nurrienrs, furrher limiring yields.
Chemical pesricides, and rhe srraregies by which
rhey are applied ro fighr crop pesrs, can have harmful
uninrended consequences. |irsr, pesrs can develop resis
rance, which means rhar higher and higher doses of pes
ricides musr be applied or new chemicals developed peri
odically ro achieve rhe same level of conrrol. Resisrance
is now found in more rhan
500 insecr and mire pesrs,
over 100 weeds, and in
abour 150 planr parhogens
(vR| 1994j. Second, popu
larions of rhe narural en
emies of pesrs are decimared
by heavy pesri ci de use.
Narural predarors are ofren
more susceprible ro synrheric
poisons rhan are rhe pesrs
because rhey have nor had
rhe same evolurionary expe
rience wirh overcoming planr
chemicals rhar rhe pesrs
rhemselves have had. And narural predarors also rypi
cally have much smaller popularion sizes rhan rhose of
rheir prey. Desrrucrion of predaror popularions leads ro
explosions in prey numbers, nor only freeing rarger pesrs
from narural conrrols bur ofren promoring" orher
nonpesr species ro pesr srarus. |n California in rhe
190s, for insrance, J4 of rhe J5 mosr imporranr agri
culrural pesrs had been elevared ro rhar srarus by rhe
overuse of pesricides (NRC 19S9j. Third, exposure ro
pesricides and herbicides may pose serious healrh risks
ro humans and many orher rypes of organisms: rhe re
cenrly discovered declines in human sperm counrs may
be arrriburable in parr ro such exposure (Colborn er al.
1996j.
|orrunarely, an esrimared 99 percenr of poren
rial crop pesrs are conrrolled by narural enemies, includ
ing many birds, spiders, parasiric wasps and flies, lady
bugs, fungi, viral diseases, and numerous orher rypes of
acriviries such as gardening
and perkeeping, narure pho
rography and filmmaking,
bird feeding and warching,
hi ki ng and campi ng,
ecorouring and mounraineer
ing, riverrafring and boar
ing, fishing and hunring, and
in a wide range of orher ac
riviries. |or many, narure is
an unparalleled source of
wondermenr and inspirarion,
peace and beaury, fulfillmenr
and rejuvenarion (e.g., Kellerr and vilsonj.
THkLATS TO LCOSYSTLM SLkvICLS
Ecosysrem services are being impaired and de
srroyed by a wide variery of human acriviries. |oremosr
among rhe immediare rhrears are rhe conrinuing desrruc
rion of narural habirars and rhe invasion of nonnarive
species rhar ofren accompanies such disruprion: in ma
rine sysrems, overfishing is a major rhrear. The mosr
irreversible of human impacrs on ecosysrems is rhe loss
of narive biodiversiry. A conservarive esrimare of rhe
rare of species loss is abour one per hour, which unforru
narely exceeds rhe rare of evolurion of new species by a
facror of 10,000 or more (vilson 19S9: Lawron and
May 1995j. bur complere exrincrion of species is only
rhe final acr in rhe process. The rare of loss of local popu
larions of speciesrhe popularions rhar generare eco
sysrem services in specific localiries and regionsis or
ders of magnirude higher (Daily and Ehrlich 1995: Hughes
er al., in prep.j. Desrroying orher life forms also disruprs
rhe web of inreracrions rhar could help us discover rhe
porenrial usefulness of specific planrs and animals (Th
ompson 1994j. Once a pollinaror or a predacious insecr
is on rhe brink of exrincrion, for insrance, ir would be
difficulr ro discover irs porenrial uriliry ro farmers.
Orher imminenr rhrears include rhe alrerarion of
rhe Earrh`s carbon, nirrogen, and orher biogeochemical
cycles rhrough rhe burning of fossil fuels and heavy use
of nirrogen ferrilizer: degradarion of farmland rhrough
unsusrainable agriculrural pracrices: squandering of fresh
warer resources: roxificarion of land and warerways: and
overharvesring of fisheries, managed foresrs, and orher
rheorerically renewable sysrems.
These rhrears ro ecosysrem services are driven
ulrimarely by rwo broad underlying forces. One is rapid,
unsusrainable growrh in rhe scale of rhe human enrer
11
Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
||gore 11Ladybug larva (Cqc|caeda pc||taj earing an aphid.
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organisms (Debach 194j.
These narural biological
conrrol agenrs save farmers
billions of dollars annually
by prorecring crops and re
ducing rhe need for chemi
cal conrrol (Naylor and
Ehrlich 199j.
Sccd D|spcrsa|
Once a seed germi
nares, rhe resulring planr is
usually roored in place for
rhe resr of irs life. |or planrs, rhen, movemenr ro new
sires beyond rhe shadow of rhe parenr is usually achieved
rhrough seed dispersal. Many seeds, such as rhose of
rhe dandelion, are dispersed by wind. Some are dispersed
by warer, rhe mosr famous being rhe seafaring coconur.
Many orher seeds have evolved ways of gerring around
by using animals as rheir dispersal agenrs. These seeds
may be packaged in sweer fruir ro reward an animal for
irs dispersal services: some of rhese seeds even require
passage rhrough rhe gur of a bird or mammal before
rhey can germinare. Orhers require burialby, say, a
forgerful jay or a squirrel which larer leaves irs cache
unearenfor evenrual germinarion. Srill orhers are
equipped wirh sricky or sharp, spiny surfaces designed
ro carch onro a passing animal and go for a long ride
before dropping or being rubbed off. virhour rhousands
of animal species acring as seed dispersers, many planrs
would fail ro reproduce successfully. |or insrance, rhe
whirebark pine (Pinus albicaulisj, a rree found in rhe
Rockies and Sierra Nevada Cascade Mounrains, cannor
reproduce successfully wirhour a bird called Clark`s Nur
cracker (Nucifraga columbianaj, which chisels pine seeds
our of rhe righrly closed cones and disperses and buries
rhem: wirhour rhis service, rhe cones do nor open far
enough ro ler rhe seeds fall our on rheir own. Animal
seed dispersers play a cenrral role in rhe srrucrure and
regenerarion of many pine foresrs (Lanner 1996j. Dis
ruprion of rhese complex services may leave large areas
of foresr devoid of seedlings and younger age classes of
rrees, and rhus unable ro recover swifrly from human
impacrs such as land clearing.
Acsthct|c 8caatq aad Iatc||cctaa| aad Sp|r|taa| St|ma
|at|ca
Many human beings have a deep appreciarion of
narural ecosysrems. Thar is apparenr in rhe arr, reli
gions, and rradirions of diverse culrures, as well as in
1J
Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
||gore 12The goods and services supplied by rhis badly deforesred and eroded region
of Madagascar are all bur gone and would be difficulr ro resrore.
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prise: in popularion size, in percapira consumprion, and
also in rhe environmenral impacrs rhar rechnologies and
insrirurions generare as rhey produce and supply rhose
consumables (Ehrlich er al. 19j. The orher underlying
driver is rhe frequenr mismarch berween shorrrerm, in
dividual economic incenrives and longrerm, socieral
wellbeing. Ecosysrem services are generally grearly un
dervalued, for a number of reasons: many are nor rraded
or valued in rhe markerplace: many serve rhe public good
rarher rhan provide direcr benefirs ro individual landown
ers: privare properry owners ofren have no way ro ben
efir financially from rhe ecosysrem services supplied ro
sociery by rheir land: and, in facr, economic subsidies
ofren encourage rhe conversion of such lands ro orher,
markervalued acriviries. Thus, people whose acriviries
disrupr ecosysrem services ofren do nor pay direcrly for
rhe cosr of rhose losr services. Moreover, sociery ofren
does nor compensare landowners and orhers who do safe
guard ecosysrem services for rhe economic benefirs rhey
lose by foregoing more lucrarive bur desrrucrive land uses.
There is a crirical need for policy measures rhar address
rhese driving forces and embed rhe value of ecosysrem
services inro decision making frameworks.
vAIUATION O| LCOSYSTLM SLkvICLS
Human sociery would cease ro exisr in rhe ab
sence of ecosysrem services. Thus, rheir immense value
ro humaniry is unquesrionable. Yer quanrifying rhe value
of ecosysrem services in specific localiries, and measur
ing rheir worrh againsr rhar of compering land uses is no
simple rask. vhen rradeoffs musr be made in rhe alloca
rion of land and orher resources ro compering human
acriviries, rhe resolurion ofren requires a measure of whar
is known as rhe marginal value. |n rhe case of ecosysrem
services, for example, rhe quesrion rhar mighr be posed
would be: by how much would rhe flow of ecosysrem
services be augmenred (or diminishedj wirh rhe preserva
rion (or desrrucrionj of rhe nexr hecrare of foresr or wer
land' Esrimarion of marginal values is complex (e.g.,
bawa and Cadgil 199: Daily 199bj. Ofren a qualira
rive comparison of relarive values is sufficienr rhar is,
which is grearer, rhe economic benefirs of a parricular
developmenr projecr or rhe benefirs supplied by rhe eco
sysrem rhar would be desrroyed, measured over a rime
period of inreresr ro people concerned abour rhe wellbeing
of rheir grandchildren'
There are, and will remain, many cases in which
ecosysrem service values are highly uncerrain. Yer rhe
pace of desrrucrion of narural ecosysrems, and rhe irre
versibiliry of mosr such desrrucrion on a rime scale of
inreresr ro humaniry, warranrs subsranrial caurion. valu
ing a narural ecosysrem, like valuing a human life, is
fraughr wirh difficulries. 1usr as socieries have recog
nized fundamenral human righrs, however, ir may be pru
denr ro esrablish fundamenral ecosysrem prorecrions even
rhough uncerrainry over economic values remains. New
insrirurions and agreemenrs ar rhe inrernarional and
subnarional level will be needed ro encourage fair parrici
parion in such prorecrions (see, e.g., Heal 1994j.
The rremendous expense and difficulry of repli
caring losr ecosysrem services is perhaps besr illusrrared
by rhe resulrs of rhe firsr biosphere J mission," in which
eighr people lived inside a J.15acre closed ecosysrem
for rwo years. The sysrem fearured agriculrural land and
replicas of several narural ecosysrems such as foresrs
and even a miniarure ocean. |n spire of an invesrmenr of
more rhan 5J00 million in rhe design, consrrucrion, and
1J
Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
operarion of rhis model
earrh, ir proved impossible
ro supply rhe marerial and
physical needs of rhe eighr
biospherians for rhe in
rended J years. Many un
pleasanr and unexpecred
problems arose, including a
drop in armospheric oxygen
concenrrarion ro 14% (rhe
level normally found ar an
elevarion of 1,500 feerj,
high spikes in carbon diox
ide concenrrarions, nirrous
oxide concenrrarions high
enough ro impair rhe brain,
an exrremely high level of
exrincrions (including 19 of
J5 verrebrare species and all
pollinarors broughr inro rhe
enclosure, which would have
ensured rhe evenrual exrinc
rion of mosr of rhe planr
species as wellj, overgrowrh
of aggressive vines and al
gal mars, and popularion
explosions of crazy anrs,
cockroaches, and karydids.
Even heroic personal efforrs
on rhe parr of rhe biospherians did nor suffice ro make
rhe sysrem viable and susrainable for eirher humans or
many nonhuman species (Cohen and Tilman 1996j.
MAJOk UNCLkTAINTILS
Sociery would clearly profir by furrher invesrigarion inro
some of rhe following broad research quesrions so rhar
we mighr avoid on biosphere 1, rhe earrh, unpleasanr
surprises like rhose rhar plagued rhe biosphere J projecr
(Holdren 1991: Cohen and Tilman 1996: Daily 199bj:
vhar is rhe relarive impacr of various human acrivi
ries upon rhe supply of ecosysrem services'
vhar is rhe relarionship berween rhe condirion of an
ecosysremrhar is, relarively prisrine or heavily
modifiedand rhe quanriry and qualiry of ecosys
rem services ir supplies'
To whar exrenr do ecosysrem services depend upon
biodiversiry ar all levels, from genes ro species ro
landscapes'
To whar exrenr have vari
ous ecosysrem services al
ready been impaired' And
how are impairmenr and risk
of furure impairmenr disrrib
ured in various regions of
rhe globe'
How inrerdependenr are
differenr ecosysrem ser
vices' How does exploiring
or damaging one influence
rhe funcrioning of orhers'
To whar exrenr, and over
whar rime scale, are ecosys
rem services amenable ro
repair or resrorarion'
How effecrively, and ar
how large a scale, can ex
isring or foreseeable human
rechnologies subsrirure for
ecosysrem services' vhar
would be rhe side effecrs of
such subsrirurions'
Civen rhe currenr srare
of rechnology and rhe scale
of rhe human enrerprise,
whar proporrion and sparial
parrern of land musr remain
relarively undisrurbed, lo
cally, regionally, and globally, ro susrain rhe delivery
of essenrial ecosysrem services'
CONCIUSIONS
The human economy depends upon rhe services performed
for free" by ecosysrems. The ecosysrem services sup
plied annually are worrh many rrillions of dollars. Eco
nomic developmenr rhar desrroys habirars and impairs
services can creare cosrs ro humaniry over rhe long rerm
rhar may grearly exceed rhe shorrrerm economic ben
efirs of rhe developmenr. These cosrs are generally hid
den from rradirional economic accounring, bur are none
rheless real and are usually borne by sociery ar large.
Tragically, a shorrrerm focus in landuse decisions ofren
sers in morion porenrially grear cosrs ro be borne by
furure generarions. This suggesrs a need for policies
rhar achieve a balance berween susraining ecosysrem
services and pursuing rhe worrhy shorrrerm goals of
economic developmenr.
||gore 13The producrion of rhis meal benefired from many
ecosysrem services, including narural pesr conrrol, pollina
rion, mainrenance of soil ferriliry, purificarion of warer, and
moderarion of climare.
14
Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
ACkNOwILD6MLNTS
ve rhank rhe Packard |oundarion and rhe Pew
|oundarion for financial supporr.
kL|LkLNCLS
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|Ok MOkL IN|OkMATION
Abcat thc Paac| c[ Sc|cat|sts
This reporr presenrs rhe consensus reached by a
panel of 11 scienrisrs chosen ro include a broad array of
experrise in rhis area. This reporr underwenr peer review
and was approved by rhe board of Edirors of |ssaes |a
Ecc|cqq. The affiliarions of rhe members of rhe panel of
scienrisrs are:
environmenr. All reporrs undergo peer review and musr
be approved by rhe edirorial board before publicarion.
LJ|tor|a| 8oarJ ot Issacs |a Fcc|cqq
Dr. David Tilman, EdirorinChief, Deparrmenr of Ecol
ogy, Evolurion and behavior, Universiry of Minnesora,
Sr. Paul, MN 5510S609. Email: rilman@lrer.umn.edu
8oarJ memhers
Dr. Srephen Carpenrer, Cenrer for Limnology, Universiry
of visconsin, Madison, v| 5J06
Dr. Deborah 1ensen, The Narure Conservancy, 1S15
Norrh Lynn Srreer, Arlingron, vA JJJ09
Dr. Simon Levin, Deparrmenr of Ecology 6 Evolurionary
biology, Princeron Universiry, Princeron, N1
0S544100J
Dr. 1ane Lubchenco, Deparrmenr of Zoology, Oregon
Srare Universiry, Corvallis, OR 9JJ1J914
Dr. 1udy L. Meyer, |nsrirure of Ecology, The Universiry of
Ceorgia, Arhens, CA J060JJJ0J
Dr. Cordon Orians, Deparrmenr of Zoology, Universiry
of vashingron, Searrle, vA 9S195
Dr. Lou Pirelka, Appalachian Environmenral Laborarory,
Cunrer Hall, |rosrburg, MD J15JJ
Dr. villiam Schlesinger, Deparrmenrs of borany and
Ceology, Duke Universiry, Durham, NC J0S
0J40
Add|t|caa| Ccp|cs
To receive addirional copies of rhis reporr or pre
vious |ssaes |a Ecc|cqq, please conracr:
Public Affairs Office
Ecological Sociery of America
J010 Massachuserrs Avenue, Nv
Suire 400
vashingron, DC J00J6
esahq@esa.org
(J0Jj SJJSJ
Special rhanks ro rhe U.S. Environmenral Prorec
rion Agency Office of Susrainable Ecosysrems and Com
muniries for supporring prinring and disrriburion of rhis
documenr.
Cover phoro credirs, clockwise from rop lefr:
Nadine Cavender, Nadine Cavender, Nadine Cavender,
Claude Cavender, 1r., Nadine Cavender, unknown, Claude
Cavender, 1r., and unknown.
Issoes |n Lco|ogy Nomher 2 Spr|ng 1997
16
Dr. Crerchen C. Daily, Panel Chair, Deparrmenr of bio
logical Sciences, Sranford Universiry, Sranford, CA 94J05
Dr. Susan Alexander, Earrh Sysrems Science and Policy,
California Srare Universiry, Monrerey bay, 100 Campus
Cenrer, Seaside, CA 9J955
Dr. Paul R. Ehrlich, Deparrmenr of biological Sciences,
Sranford Universiry, Sranford, CA 94J05
Dr. Larry Coulder, Deparrmenr of Economics, Sranford
Universiry, Sranford, CA 94J05
Dr. 1ane Lubchenco, Deparrmenr of Zoology, Oregon
Srare Universiry, Corvallis, OR 9JJ1
Dr. Pamela A. Marson, Environmenral Science Policy and
Managemenr, Universiry of California, berkeley, CA
94J0
Dr. Harold A. Mooney, Deparrmenr of biological Sciences,
Sranford Universiry, Sranford, CA 94J05
Dr. Sandra Posrel, Clobal varer Policy Projecr, 10 Lark
spur Drive, Amhersr, MA 0100J
Dr. Srephen H. Schneider, Deparrmenr of biological Sci
ences, Sranford Universiry, Sranford, CA 94J05
Dr. David Tilman, Deparrmenr of Ecology, Evolurion and
behavior, Universiry of Minnesora, Sr. Paul, MN 5510S
609
Dr. Ceorge M. voodwell, voods Hole Research Cenrer,
P.O. box J96, voods Hole, MA 0J54J
Much of rhe informarion in rhis reporr was de
rived from C. Daily, ediror. 199. Narure`s Services:
Socieral Dependence onNarural Ecosysrems. |sland
Press, vashingron, D.C.
Abcat thc Sc|cacc Wr|tcr
Yvonne baskin, a science wrirer, edired rhe re
porr of rhe panel of scienrisrs ro allow ir ro more effec
rively communicare irs findings wirh nonscienrisrs.
Abcat Issacs |a Fcc|cqq
|ssaes |a Ecc|cqq is designed ro reporr, in lan
guage undersrandable by nonscienrisrs, rhe consensus
of a panel of scienrific experrs on issues relevanr ro rhe

Abcat Issacs |a Fcc|cqq


|ssaes |a Ecc|cqq is designed ro reporr, in language undersrandable by nonscienrisrs, rhe
consensus of a panel of scienrific experrs on issues relevanr ro rhe environmenr. |ssaes |a
Ecc|cqq is supporred by rhe Pew Scholars in Conservarion biology program and by rhe Eco
logical Sociery of America. |r is published ar irregular inrervals, as reporrs are complered. All
reporrs undergo peer review and musr be approved by rhe Edirorial board before publicarion.
|ssaes |a Ecc|cqq is an official publicarion of rhe Ecological Sociery of America, rhe narion`s
leading professional sociery of ecologisrs. |ounded in 1915, ESA seeks ro promore rhe
responsible applicarion of ecological principles ro rhe solurion of environmenral problems.
|or more informarion, conracr rhe Ecological Sociery of America, J010 Massachuserrs Av
enue, Nv, Suire 400, vashingron, DC, J00J6. |SSN 109JS9S

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