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Benefits of Plant Diversity to Ecosystems: Immediate, Filter and Founder Effects Author(s): J. P. Grime Source: Journal of Ecology, Vol.

86, No. 6 (Dec., 1998), pp. 902-910 Published by: British Ecological Society Stable URL: . Accessed: 04/03/2014 15:37
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Journal of Ecology1998, 86,902-910



Benefits ofplantdiversity to ecosystems: immediate, filter and founder effects

Un1it of Comparative ofAnimal Plans1t aniid Plans1t Scienices, ofSheffield, Ecology,Departmenit University SJO2TN, UK Sheffield

1 It is usefulto distinguish betweenthe immediateeffects of species richnesson and thosewhichbecomeapparenton a longertimescale, described ecosystems here as filter and founder effects. 2 Relationships between plantdiversity and ecosystem can be exploredby properties - dominants, classifying componentspeciesinto threecategories and subordinates in particular transients. Dominantsrecur are relatively exhibit vegetation types, large, forresources coarse-grained foraging and, as individualspecies,make a substantial to theplantbiomass.Subordinates also showhighfidelity ofassociation contribution withparticular vegetation typesbut theyare smallerin stature, forageon a more restricted scale and tendto occupymicrohabitats delimited and by the architecture oftheir a heterogeneous phenology associateddominants. Transients comprise assortmentof speciesof low abundanceand persistence; a highproportion arejuvenilesof in neighbouring or subordinates speciesthatoccuras dominants ecosystems. 3 A 'mass ratio'theory controls are in proportion to inputs proposesthatimmediate to primary are determined to an overwhelming extent production, by the traitsand functional to therichness ofthedominant insensitive diversity plantsand arerelatively of subordinates and transients. Recentexperiments themassratiohypothesis support and theconclusion of Huston(1997) thatclaimsofimmediate ofhighspecies benefits to ecosystem richness functions arisefrom of data. misinterpretation 4 Attribution of immediatecontrolto dominantsdoes not exclude subordinates and transients in the determination frominvolvement of ecosystem and function ifintermittent, Bothare suspected to play a crucial, sustainability. rolebyinfluencing the recruitment of dominants.Some subordinates may act as a filter influencing regeneration by dominants following majorperturbations. 5 Transients from theseed rainand seed banksand providean indexof the originate and subordinates at specific sites.Wherethe landscape pool of potentialdominants in the reservoir carousel operates against a backgroundof decliningdiversity of loss of ecosystem we may predict thata progressive functions colonizingtransients, will arise fromthe declinein the precision withwhichdominants can engagein the of ecosystems. and relocation re-assembly
Keywords. biodiversity, dominance,ecosystem function, landscape ecology,regen-


Journal of Ecology (1998)

86, 902-9 10
organismsare eliminatedor debilitated, it is often possibleto demonstrate a causal connection between losses in biodiversity and declinesin ecosystem function and in benefits to humans(Smith1968; LeCren etal. 1972; Pearson & Rosenberg1976; Vitousek& Melillo 1979). A more difficult subjectfor analysis arisesin circumstances whereeither ecospecies-poor

When ecosystems become degradedby pollutionor over-exploitation to a pointwhere formerly dominant Correspondence: J. P. Grime (e-mail j.p.grime((7)


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903 J.P. Grinme

systems (e.g. boreal forests, bogs and heathland)or species-rich ecosystems(e.g. limestonegrasslands, tropicalforests and coral reefs)remainin existence but experiencegradual losses in species or genetic diversity (Thomas 1960; Bobbink & Willems 1987; Barr etal. 1993). Does such attrition, particularly whereit affects speciesof low relative abundancein communities, have major implications forecosystem function and viability? To addressthisquestionit is helpful to recognize two separateissues. First,we need to know whether losses in speciesrichness have immediate (proximal) effects on ecosystem function. itis necessary Secondly, to considerthe possibility of less conspicuouslongon termconsequences.This paper comments briefly bothissues.Evidenceis drawnmainly from studies of herbaceousvegetation but thereappear to be some principles thatapplymorewidely.

Dominants, subordinates and transients

In orderto estimate ofa speciesloss theconsequences it is necessary to knowwhat upon itshostecosystem, it role (if any) the organism concernedplays within (Grime 1973; Whittaker1975; McNaughton 1978; Lawton 1994). In his pioneering to define attempts thefunctional rolesofspecieswithin plantand animal communities, Whittaker (1965, 1975)recognized that a useful firststep is to order componentspecies to their relative according abundanceor productivity. When large numbersof the resulting 'dominanceit is possible to diversity' profilesare constructed associationsbetween beginthe searchforconsistent thetraits ofspeciesand their abundanceinecosystems and communities. For European herbaceousvegetation, thereis an on theabundanceand enormous fundofinformation characteristics of component species in relatively small(c. 1M2) vegetation samples.Discussionsof the of thesedata are available in functional significance Grime(1973, 1987),Grubbetal. (1982) and Mitchley & Grubb (1986). Many ecological factorsand plant deserve consideration as potential determinants traits of dominance-diversity and it is clear that profiles, siteto site.Howeffects controlling varyindetailfrom can be attempted and this is ever,a generalization in Fig.1 in theform ofan idealiseddomisummarized
nance-diversity curve

smaller instature (Fig. 2) and form a lowerproportion the In contrast to the suborof biomass. marked and lack dinates,the transients are heterogeneous fidelity of association with particulardominants. to the They make a very small total contribution and in functional traits vegetation and varyinnumber onlyas seedto a greatextent. Most are represented are species lingsor juvenilesand a highproportion in other ecoor subordinates thatoccuras dominants logical situations (Table 1) oftensituatednearby.In to notethatformal procedures passing, itis interesting used to collectand analysedata on the speciescom1905; positionof plant communities (e.g. Clemnents Braun-Blanquet1932; Bray & Curtis 1957; Kent & of under-recording Coker 1992) often have theeffect or discarding information on transients which,from as a classificatory are frequently regarded viewpoint, 'misfits'. byspecies Table 1 showsthatsuchexcursions appear commonplacewhen detailedand exhaustive samplingproceduresare applied: recordsfromall threeof the sampled habitatsinclude many species habitats. thatare moretypically associatedwithother

ofbiodiversity Immediate effects

Can we generalize about the relative of importance dominants,subordinatesand transientsas determinants ofecosystem suchas productivity, properties relations, nutrient cycling carbonsequestration, water and storage, litter and resistance and resilience quality Both theory and experimental evito perturbations? dence (Huston 1997; Aarssen 1997) suggestthatthe funcextent to whicha plantspeciesaffects ecosystem tionsis likelyto be closelypredictable fromits conto thetotalplantbiomass.This 'mass ratio' tribution and hypothesisis implicitin many commentaries to ecosystem function modelsrelating (Shugart1984; Pastor & Post 1986; Huston & Smith 1987; Grime 1987; Sala etal. 1996; Huston 1997) and is dictated and chemistry thatrequire that bythelaws ofphysics within involve of autotrophs ecosystems largeeffects in syntheses, and in inputs to major participation resource fluxes and degradative processes.It follows to a thatecosystem shouldbe determined properties of the dominants large extentby the characteristics in species insensitive to variation and willbe relatively in circumstances to richness wherethisis attributable in thenumber ofsubordinates and transients. changes to specify It is important thatthemass ratiohypothin applicationto the role of autoesis is restricted trophs in ecosystemprocesses. When attentionis turnedto othertrophicelements, such as parasites, the possibility herbivores, predatorsand symbionts, arisesforecosystem impactsthatare less predictably relatedto abundance. A growing bodyofexperimental evidence supports are strongly thehypothesis thatecosystem properties influenced by thecharacteristics of dominant plants. In a comparative study oftheresistance and resilience

Whittaker 1965). This

themajority of herbaceousvegthatwithin suggests etation samples threeelementscan be recognized, in speciesrichness each capable of varying and taxoas dominants, subnomicidentity, and heredescribed ordinatesand transients. The dominants are usually few in number,taller and more expansivein morphology and account for a high proportionof the ? 1998 British1 Ecological Society, biomass (as seen fora grasslandexamplein Fig.2). Many subordinates consistently co-occur withparticular dominants and,although they areusuallym1ore numerous as individuals thanthedominlants, they are

Jounl( ?1 of Ecology,


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D D Dominant S Subordinate T Transient

Benefits ofplan1t
diver-sit to





60 50 40 30 20 10 0 5 S'S- S-S-TIT S-TIS-TT-T-T-TI T TT-T 10 15 20 25


D D\

Species ranking The curve (senisiu Whittaker1965) for a small sample of herbaceousvegetation. Fig. 1 An idealised dominance-diversity value distinction is based upon relativeabundance and has been set at an arbitrary betweendominantsand subordinates and persistin the vegetation (10%). Transientsare distinguished fromsubordinates by theirfailureto regenerate under scrutiny.

C9 1998British Ecological Society, ofEcology, Journ-iial 86, 902-910

of five herbaceouscommunities to drought, late frost et al. 1995),predictions and fire (MacGillivray using traitsmeasuredin the laboratory were foundto be accuratewhencalculationswere based upon means to therelative weighted according abundanceofeach conplantspeciesat each experimental site.A similar clusion was drawn fromtwo recentinvestigations (Wardle etal. 1997; Hooper & Vitousek 1997) in whichvariousecosystem werefoundto be properties withthefunctional strongly correlated characteristics of thedominant to thebiomass. contributors There is also strong evidence that functional betweenco-existing dominantscan have differences profoundeffects on ecosystems, particularly in susis fluctuation there taining yieldoverperiodsin which in climate or vegetationmanagement.From both and experiments monitoring studies (Willis1963;Mellinger & McNaughton1975;Kemp & Williams1980; Grime etal. 1985) thereis abundant evidencethat inphenology, differences occurbetween codominants, photosynthetic mechanism, rooting depthand reprocases (Spedding& ductivebiology.In some reported & Eadie 1973; Hooper Diekmahns 1972; Armstrong & Vitousek 1997) such complementary exploitation has been shownto confer a benefit to productivity. If the immediateinfluence of vegetationon the properties of ecosystems is determined primarily by it is necessary to consider traitsof the dominants, what additional effects mightbe exertedby suband transients. ordinates First,perhaps,it shouldbe notedthatthereis no a priorireasonto suspectthat must influence such minor contributors ecosystem

their reflect thefact functioning; presence maysimply thatconditions in thepast or present have prevailing allowedthemto be admitted. However,severalsources of evidencesuggest thatsome subordinate membersofplantcommunities (as distinct from transients) fulfil rolesthatextend beyond thatofmereadventives. We may suspectthattheconsistent associationsevident between certain dominants and their subordinates (e.g. Fig. 2) reflect a complementary exploitationofhabitatresulting in a morecomplete capture of resourcesand minorbenefits to productivity. In somepairings ofsubordinates with dominants, spatial and temporal interlocking finds precise morphological expression, as in the case of the bryophyte in winter, which, Brachytheciumn rutiabuluin colonizes of the tall herb Urtica each fallenstemof the litter dioica (Furness & Grime 1982). More oftencomplementarity between dominants and subordinates in whichthe latterexploit consistsof circumstances unfavourable These maybe relatively microhabitats. as inthecase ofthecreeping herbs expressed spatially, and bryophytes thatoccupythe shaded lowerstrata of herbaceouscanopies (Al-Mufti etal. 1977;Fig. 2), or temporally, as exemplified by the small winter annualsand spring thatmakea minor geophytes conto the biomass of species-rich tribution calcareous grasslands (Ratcliffe 1961;Grimeetal. 1985). of mechanisms Further evidence ofhabitatexploitationcomplementarity betweendominantand subordinatemembers of a plantcommunity is available froman experiment (Campbell etal. 1991) in which the abundance of eightplant species in an exper-

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(a) U Festuca ovina B Agrostisspp. o Carex flacca O Carex panicea

J.P. Grimi1e
15-19.99cm 10-14.99cm
5-9.99cm _


2-2.99cm 1-1.99cm <1cm 0










Number ofcontacts (b) Carexcaryophyllea rotundifolia *Viola riviniana o Carexpulicaris M Campanula

0, ._


10-14.99cm 5-9.99cm 44.99cm 3-3.99cm 2-2.99cm 1-1.99cm <1cm

0 50 100

ofcontacts Number of the leaf canopy of (a) fourselecteddominantand (b) fourselectedsubordinate Fig. 2 Verticaldistribution component in Junewas estimated UK. Canopy distribution species in an ancientlimestone pastureat Buxton,North Derbyshire, by contactswith375 randomly vertical measuring distributed, pins(S. H. Hillier, unpublished data).

in threedistinctive Table 1 Ecological classification of thespeciesrecorded contrasted and highly habitatssampledwidelyin an area of 2400 km2around Sheffield, in terms UK. All thespeciesencountered within a particular habitatwereclassified of their habitat(columns3-6). Details of thesampling, and habitatclassification are providedin primary recording procedures
Grime etal. (1988)


Numberof Woodland m2samples species 51 40 55 65 7 5

Grassland species 23 64 38

Arable species 1 8 69

Others* 5 0 2

Total 94 79 114

Ecological Society, ofEcologyv, Journacl 86, 902-910

?) 1998British

Woodland on limestone Meadows Cereal arable

* Includesspecies associatedwith wetland or skeletal habitats (cliffs, wallsand rockoutcrops). primnarily

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906 o plant Beniefits to diversity ecosystems

imental mixture was accurately predicted from indeof shoot and root foraging pendentmeasurements in a standardized by isolatedplantsgrowing patchy environment. From this investigation it was concluded that dominancewas achieved by the develin which opment of a coarse-grained architecture mainrootsand shootsspreadrapidly through a large with rather concentration volumeofhabitat imprecise in resource-rich sectors.A complementary foraging mechanismwas recognized in subordinates;here resource capturewas achievedby a precisebut local concentration of roots and shoots in resource-rich patches,a specializationlikelyto carrythe penalty of subordination and ultimately of (in circumstances and consolidationby the domiunrestricted growth nants)riskof competitive exclusion. We mayconclude, thatfunctional divertherefore, and perhapsalso within subsityamong dominants ordinatesis capable of immediateimpacts on the of ecosystems. properties However,it may be a misof take to narrow the search for beneficial effects of immediate to an examination biodiversity effects; theroleofminor vegetation components maybecome evident only when viewed in the long term. The remainderof this paper outlinestwo hypothetical mechanismswherebybenefitsof biodiversity may ofminor intermittent effects contributors accruefrom to the plant biomass. The first involves hypothesis thesecondtransients. subordinates,

Filter effectsof biodiversity; a role for subordinates?

So farin thispaper it has been convenient to regard theplantcommunity as a stablehierarchy containing In reality, of subordinates and transients. dominants, in comfluctuations course,communities experience positiondriven by seasonal and longer-term changes in climate, and vegetation and herbivory management of component by theintrinsic dynamics plantpopufield lations.Moreover, from observations and experithere is strong circumstantial evidence thatthe ments of subordinatesin both grassland and persistence woodland vegetation is frequently dependentupon periodicevents(e.g. droughts, frosts, floods,windthrows, grazing, trampling, burning, coppicing)that thevigourand competitive effects restrict temporarily ofdominant contains plants.The literature manyrefwhereabatementof such erencesto circumstances has led to expansions events damaging bydominants, losses of subordinates and a rapid declinein species richness(Tansley & Adamson 1925; Thomas 1960; Smith etal. 1971).Therecan be little doubt,therefore, thatplantspeciesthathabitually dominate particular plantcommunities usuallyexert controlling effects on thefitness of theirsubordinates. However,sinceit is suspectedthatthe immediate controlson ecosystem properties are largely determined by thedominants it is muchmorerelevant to thepurposesofthispaperto

g 1998British Ecological Society, Jouti7ial ofEcology, 86, 902-910

ask 'Do subordinate of plant communities members exercise controls on theidentity, functional diversity and relative abundanceof dominants?' In order to review the opportunities for subordinates to control dominants it is necessary to considerthe long-term and the dynamicsof vegetation regenerative phases in the lifecyclesof dominants. Fromstudies ofvegetation succession (e.g. Watt1925, 1947) it is established thatcontinueddominanceby determined particular speciesis frequently bythesuccess of seedlingor vegetative re-establishment following disturbanceevents causing mortalitiesof a local or catastrophic dominants on either scale.Here it maybe important to recognize thatoften theearly a disturbance is a temcourse of eventsfollowing porary expansion in the cover and vigour of subThisphenomenon is mostobviousin forest ordinates. where a denselowcoverofshrubs, herbs and clearings forregenerating bryophytes mayprovidethecontext trees (Watt 1925;Skutch1929;Marks 1974;Bormnann & Likens 1979), but similarphenomenahave been described for grasslands and heathlands (Oosting of seedling 1942;Keever 1950;Hillier1990).Patterns establishment and vegetative disturbances following are not determined traits exclusively by regenerative such as the size and number of propagulesand their dispersal,dormancy,morphologyand physiology. with They arise also throughcomplex interactions in whichcontributions conditions to the substratum groundcoverby subordinate plantsmaybe expected to have both positiveand negativeeffects (Cavers & Harper 1967; Ross & Harper 1972; Grubb 1977; Connell& Slatyer 1977;Noble & Slatyer 1979;Pickett & White1985;Bazzaz 1986;Maguire& Forman1983; Burke& Grime1996).Benefits to establishment have in circumstances been described whereseedlings survive in the shelter afforded by low-growing shrubs, herbs and bryophytes (Lawrence & Hulbert 1960; Ward 1990; Hillier 1990). Negativeeffects of shrub, and bryophyte coveron theestablishment herbaceous ofgrassland and forest dominants havebeenobserved (Wardle 1959;Niering& Goodwin 1962;Webb etal. 1972; Pons 1989) and it is widelyaccepted (Fenner treesand shrubs 1992) thatmanysmall-seeded herbs, are incapableofestablishment wherethere is a closed coverofvegetation. Thereis someevidence thatquite inconspicuous subordinatemembers of the plant can exerta selectiveeffect on seedling community dominants. For example, populationsofregenerating in a microcosm in Grime(1987) experiment reported an algal film on thesoil surface thedevelsuppressed estabopmentof small-seeded speciesbut permitted lishment of larger-seeded of grasses.The significance subordinatesin plant communities therefore may extendbeyond any immediatecontributions to the carbon economy and nutrient dynamics.Over the longertermthereappears to be a potentialforsubordinatemembers of a plant community to act as a filter selecting between different potential dominants

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907 J.P. Grimle

a during theearlyphases of recolonization following disturbance event. Such selectioncould operate on thebasisofvariation intheseedreserves ofdominants and theassociateddifferences in thecapacityof their seedlings to penetrate a low canopy(Grime& Jeffrey 1965; Westobyetal. 1992). Alternatively, according to the characteristics of the groundcover the filter upon might discriminate between dominants thatrely by perrapid emergence and thosewhichregenerate sistent juveniles(Marshall 1927; Chippindale 1932; Marks 1974). Controllingeffectsof subordinates dominants upon regenerating mayalso occurthrough moreindirect mechanisms, such as provisionof sites in whichseed predation is reduced(Thompson 1987; van Tooren 1988). In this reviewit has been consideredprudent to restrict discussionof the possible role of subordinates to ratherdirecteffects filtering on therecruitment It wouldbe possible, ofdominants. to includecases wherethe impactof subhowever, ordinates arisesfrom morecomplexphenomena such of critical heras the maintenance pests,pathogens, bivoresor mutualists (e.g. Gilbert 1977; Huston & Gilbert1996). role for subordinates during Evidence of a filter remains ecosystem re-assembly anecdotal.Thereis an need forcarefully experiurgent designed, long-term ments (e.g. Ward 1990)to evaluatethisphenomenon.

a role fortranFoundereffects of biodiversity; sients?

transients On first inspection, appear to be irrelevant indito ecosystem function. They occur as scattered as seedlings viduals and many appear only briefly that fail to survive.Familiarexamplesin European forexample, includeannualssuchas specgrasslands, ies of Papaverand Polygoinan thatoccuras a legacy of former arable cultivations or seedlingsof windblownor bird-dispersed herbs,shrubsand trees.Can of vegsuch minor and incongruousconstituents and viabilityof ecoetation affect the functioning systems? In orderto explorethe ecological significance of it is usefulto identify the originof these transients individualsand to considerwhy some communities contain a wider diversity than others.This review of transients concentrates on thepossiblesignificance withwhichpotenas an indicator of theeffectiveness tialdominants are dispersed acrossthelandscapeand into 'suitable' ecosystems. However, it is recruited thattransients a neglected also worth noting represent of studies subjectin plant ecology and are worthy there beyondthe scope of thisreview.In particular, thattheaccumulation is a need to testthehypothesis in species-rich and persistence oftransients vegetation is an indicationthat a low intensity of competition prevailsin suchconditions. The sources of thletransienlts appear to be seed the surbanks in the soil and the seed rain fromn

rounding landscape.This suggests thatthetransients mayprovideusefulinformation concerning thepool of potentialcolonizingspeciesat each site.We may predict thata diversity oftransients signifies thepresof colonizersand a high ence of a rich assortment probability that,in the eventof habitatdisturbance or changes in management, therewill be a rapid ingressof different plant functional types,some of which may be capable of exploiting the new conditions. Here a specific examplewouldbe thebenwherean abandoned efitto woodland development grasslandalready contains a diverseassortment of treeseedlings. in Europe and in Efforts to conservebiodiversity manyotherpartsof theworldtakesplace in a fragmenting landscapemosaic continuously disturbed by natural events and by urbanization,arable cultiand variousforms manof grassland vation,forestry conservation therefore agement.Successful depends in part upon continuousmovement of populations and re-assembly of vegetation typesand ecosystems. The extent to whichcommunities and ecosystems are is likely to be relatedto theresrapidly reconstituted of colonizers, ervoir manyofwhichshouldbe detectas transient able priorto disturbance constituents of the existing vegetation.Following Egler (1954) we with may suspectthat the speed and completeness which ecosystem re-assembly occurswilldependupon and subdominants earlycolonization byappropriate latearrival to delayestabordinates; maybe expected lishment of a species and may even exclude some completely (Keever 1950; Niering& Goodwin 1962; Holt 1972;Platt 1975). It follows, thatthe therefore, in manycontemporary declinein diversity occurring a consequence situ losses ofini landscapesis notsimply within communities. Impoverishment mayalso occur in theprocesses a progressive ofplant failure through this failure dispersal and ecosystemre-assembly; should be detectableas a decline throughtime in in plant and speciesrichness of transients thedensity communities.

forecologicaltheory Implications
This paper has soughtto connectrecentstudiesof and ecosystem function plant diversity (e.g. Lawton 1994; Naeem etal. 1994; Tilman & Downing 1994) with another literature concernedwith the mechanismscontrolling itself. Reference to invesdiversity tigations such as those of Whittaker (1975), McNaughton (1978) and Grime (1973) revealsthat even in species-rich vegetationmost of the plant biomass may residein a small numberof dominant ofwhichare likely to overspecies,thecharacteristics ride as ecosystemcontrollersthe effects of more subordinate or transient components. As nu1nerous discussedby Huston (1997), thissuggests a need for cautious assessmentof studies (Naeem eta!l. 1994; & Downing 1994; Tilman Tilman Tilmnan eta!l. 1996;

2C 1998British Ecological Society, ofEcology, Joutrnoal 86, 902-910

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908 Benefitsofplant to diver-sity ecosystenms

etal. 1997) in which correlationsare established but and ecosystem properties between speciesrichness with respectto the relative data are not presented abundance of component species. Extreme care differences inspeciesrichness appearsnecessary where in ecosystem withdifferences are not onlycorrelated withcontrasting functions but are also confounded and resourcedynamics(e.g. Tilman & lifehistories Downing 1994). However,untilthe resultsof longthe functional charactermexperiments measuring ofconteristics ofnatural and synthesized ecosystems composition and functional trastedspecies richness of of immediate effects are available, the possibility speciesrichness perise cannotbe eliminated. Even if the balance of evidence (Huston 1997; themass ratio Grime1997)continues to shift towards hypothesis and against the propositionthat species of ecofunctioning richness controlsthe immediate systems, thisdoes notmean thatlossesofplantdiversity should be viewed with equanimity.Declining diversity maybe associatedwithless obviousimpacts and founder that operate throughfailuresin filter In particular effects. we suspectthattheremay be a in circumstances where loss of functions progressive vegetation patch dynamics and ecosystem rethebackground ofa declincontinue against assembly to this ingpool of colonizingpropagules.According of plant diversityin hypothesisthe significance functions may of ecosystem relationto deterioration of arise primarily its effects from on the recruitment of speceffects dominants rather thananyimmediate ies richness pe- se.

I am grateful to Sue Hillier,JohnHodgson and Ken to use data obtained in Thompson for permission and to Sarah Buckland,Andcollaborative projects, rew Askew and Suzanne Hubbard for assistancein It is a pleasure to preparationof the manuscript. comand constructive acknowledgethe perceptive mentsof Sandra Lavorel and Michael Huston on an ofthispaper.Some oftheresearch drawn earlier draft upon in this paper was supportedby the Natural Environment ResearchCouncil.

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J.P. Grimce

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