Supplied ro Human Socieries by Narural Ecosysrems l s s u : s
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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. P h o r o
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S e r v i c e J 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. P h o r o
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R i c k e r r s 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 P h o r o
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R i c k e r r s 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 P h o r o
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R i c k e r r s ||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. P h o r o
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E h r l i c h 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. P h o r o
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S e r v i c e ||gore 7Herbal pharmacisr in Dali, Yunnan Province, China. An esrimared S0 percenr of rhe world`s popu larion relies on narural medicinal producrs. P h o r o
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S e r v i c e 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. P h o r o
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D a i l y 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. P h o r o
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S e r v i c e 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. P h o r o
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S e r v i c e 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. P h o r o
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S e r v i c e 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. P h o r o
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D a i l y 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 Adams, R. McC. 19S1. Hearrland of Ciries: Surveys of Ancienr Serrlemenr and Land Use on rhe Cenrral |loodplain of rhe Euphrares, Chicago: Universiry of Chicago Press. Alexander, S., S. Schneider, and K. Lagerquisr. 199. Ecosysrem services:|nreracrion of Climare and Life. Pages 19J in C. Daily, ediror. Narure`s Services: Socieral Dependence on Narural Ecosysrems. |sland Press, vashingron, D.C. Allegre, C. and S. Schneider. 1994. The evolurion of rhe earrh. 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Earrhworms: Their Ecology and Relarionships wirh Soils and Land Use. Academic Press, New York. Leung, A.Y. and S. |osrer. 1996. Encyclopedia of Common Narural |ngredienrs Used in |ood, Drugs, and Cosmerics. 1ohn viley 6 Sons, |nc., New York. McCormick, K.D., M.A. Deyrup, E.S. Menges, S.R. vallace, 1. Meinwald, and T. Eisner. 199J. Relevance of chemisrry ro conservarion of isolared popularions: rhe case of volarile leaf componenrs of Dicerandra minrs. Proc. Nar. Acad. Sci. USA 90: 0105. 15 Issoes |n Lco|ogy Nomher 2 Spr|ng 1997 Myers, N. 19SJ. A vealrh of vild Species. vesrview Press, boul der, CO. Myers, N. 199. The worlds foresrs and rheir ecosysrem services. Pages J15JJ5 in C. Daily, ediror. Narure`s Services: Socieral Dependence on Narural Eco sysrems. |sland Press, vashingron, D.C. Nabhan, C.P. and S.L. buchmann. 199. Pollinarion services: biodiversiry`s direcr link ro world food srabiliry. Pages 1JJ150 in C. Daily, ediror. Narure`s Services: Socieral Dependence on Narural Ecosysrems. |sland Press, vash ingron, D.C. Narional Research Council (NRCj. 19S9. Alrernarive Agriculrure. Narional Academy Press, vashingron, D.C. Narional Research Council (NRCj. 199J. Managing Clobal Ceneric Resources: The U.S. Narional Planr Cermplasm Sysrem. Narional Academy Press, vashingron, D.C. Naylor, R. and P. Ehrlich. The value of narural pesr conrrol services in agriculrure. Pages 15114 in C. Daily, ediror. Narure`s Services: Socieral Dependence on Narural Ecosysrems. |s land Press, vashingron, D.C. Oldeman, L., v. van Engelen, and 1. Pulles. 1990. The exrenr of humaninduced soil degradarion, Annex 5" of L. R. Oldeman, R. T. A. Hakkeling, and v. C. Sombroek, vorld Map of rhe Srarus of Human|nduced Soil Degradarion: An Explanarory Nore, rev. Jd ed. vageningen: |nrernarional Soil Reference and |nformarion Cenrre. OvergaardNielsen, C. 1955. Srudies on enchyrraeidae J: |ield srud ies. Narura 1urlandica 4: 55S. 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The foresr and rhe hydrological cycle. Pages JJJ94 in R. Dickinson, ediror. The Ceophysiology of Amazonia. 1ohn viley and Sons, New York. Schlesinger, v. 1991. biogeochemisrry: An Analysis of Clobal Change. Academic Press, San Diego. Schneider, S. and P. bosron, edirors. 1991. Scienrisrs on Caia. M|T Press, bosron. Schneider, S. and R. Londer. 19S4. The Coevolurion of Climare and Life. Sierra Club books, San |rancisco. Shiklomanov, |.A. 199J. vorld fresh warer resources. Pp. 1JJ4 in P. Cleick, ediror. varer in Crisis: A Cuide ro rhe vorld`s |resh varer Resources. Oxford Universiry Press, New York. Shukla, 1., C. Nobre, and P. Sellers. 1990. Amazon deforesrarion and climare change. Science. J4: 1JJJ1JJ5. Tilman, D. 199. biodiversiry and ecosysrem funcrioning. Pages 9J11J in C. Daily, ediror. Narure`s Services: Socieral Dependence on Narural Ecosysrems. |sland Press, vash ingron, D.C. Thompson, 1.N. 1994. The Coevolurionary Process. Chicago Univ. Press, Chicago. Unired Narions |ood and Agriculrure Or ganizarion (UN|AOj. 199J. Marine |isheries and rhe Law of rhe Sea: A Decade of Change. |isheries Circular No. S5J, Rome. Unired Narions |ood and Agriculrure Organizarion (UN|AOj. 1994. |AO Yearbook of |ishery Srarisrics. volume 1. Unired Srares Deparrmenr of Agriculrure (USDAj. 199J. vorld Ag riculrure: Trends and |ndicarors, 19091. vashingron, DC: USDA. virousek, P., P. Ehrlich, A. Ehrlich, and P. Marson. 19S6. Human appropriarion of rhe producrs of phorosynrhesis. bioScience J6: J6SJJ. virousek, P., 1. Aber, R. Howarrh, C. Likens, P. Marson, D. Schindler, v. Schlesinger, and D. Tilman. 199. Human alrerarion of rhe global nirrogen cycle: causes and consequences. [COM PLETE] vilson, E.O. 19S. The lirrle rhings rhar run rhe world: The impor rance and conservarion of inverrebrares. Conservarion bi ology 1: J44J46. vilson, E.O. 19S9. Threars ro biodiversiry. Scienrific American Sepr: 10S116. vorld bank. 1991. Agriculrural biorechnology: The Nexr Creen Revolurion' vorld bank Technical Paper no. 1JJ. vash ingron, D.C. vorld Resources |nsrirure (vR|j. 1994. vorld Resources: A Cuide ro rhe Clobal Environmenr. Oxford Universiry Press, Ox ford. |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