Journal of

Ecology 0887\
75\ 891809

ESSAY REVIEW

Bene_ts of plant diversity to ecosystems] immediate\

_lter and founder effects
J[P[ GRIME
Unit of Comparative Plant Ecology\ Department of Animal and Plant Sciences\ University of Shef_eld\
Shef_eld S09 1TN\ UK

Summary
0 It is useful to distinguish between the immediate e}ects of species richness on
ecosystems and those which become apparent on a longer time scale\ described here
as _lter and founder e}ects[
1 Relationships between plant diversity and ecosystem properties can be explored by
classifying component species into three categories dominants\ subordinates and
transients[ Dominants recur in particular vegetation types\ are relatively large\ exhibit
coarse!grained foraging for resources and\ as individual species\ make a substantial
contribution to the plant biomass[ Subordinates also show high _delity of association
with particular vegetation types but they are smaller in stature\ forage on a more
restricted scale and tend to occupy microhabitats delimited by the architecture and
phenology of their associated dominants[ Transients comprise a heterogeneous assort!
ment of species of low abundance and persistence^ a high proportion are juveniles of
species that occur as dominants or subordinates in neighbouring ecosystems[
2 A {mass ratio| theory proposes that immediate controls are in proportion to inputs
to primary production\ are determined to an overwhelming extent by the traits and
functional diversity of the dominant plants and are relatively insensitive to the richness
of subordinates and transients[ Recent experiments support the mass ratio hypothesis
and the conclusion of Huston "0886# that claims of immediate bene_ts of high species
richness to ecosystem functions arise from misinterpretation of data[
3 Attribution of immediate control to dominants does not exclude subordinates
and transients from involvement in the determination of ecosystem function and
sustainability[ Both are suspected to play a crucial\ if intermittent\ role by in~uencing
the recruitment of dominants[ Some subordinates may act as a _lter in~uencing
regeneration by dominants following major perturbations[
4 Transients originate from the seed rain and seed banks and provide an index of the
pool of potential dominants and subordinates at speci_c sites[ Where the landscape
carousel operates against a background of declining diversity in the reservoir of
colonizing transients\ we may predict that a progressive loss of ecosystem functions
will arise from the decline in the precision with which dominants can engage in the
re!assembly and relocation of ecosystems[
Keywords] biodiversity\ dominance\ ecosystem function\ landscape ecology\ regen!
eration
Journal of Ecology "0887# 75\ 891809
Introduction
When ecosystems become degraded by pollution or
over!exploitation to a point where formerly dominant

891

Correspondence]
she.eld[ac[uk#[

J[

P[

Grime

"e!mail

j[p[grime

organisms are eliminated or debilitated\ it is often

possible to demonstrate a causal connection between
losses in biodiversity and declines in ecosystem func!
tion and in bene_ts to humans "Smith 0857^ LeCren
et al[ 0861^ Pearson + Rosenberg 0865^ Vitousek +
Melillo 0868#[ A more di.cult subject for analysis
arises in circumstances where either species!poor eco!

892
J[P[ Grime

systems "e[g[ boreal forests\ bogs and heathland# or

species!rich ecosystems "e[g[ limestone grasslands\
tropical forests and coral reefs# remain in existence
but experience gradual losses in species or genetic
diversity "Thomas 0859^ Bobbink + Willems 0876^
Barr et al[ 0882#[ Does such attrition\ particularly
where it a}ects species of low relative abundance in
communities\ have major implications for ecosystem
function and viability<
To address this question it is helpful to recognize
two separate issues[ First\ we need to know whether
losses in species richness have immediate "proximal#
e}ects on ecosystem function[ Secondly\ it is necessary
to consider the possibility of less conspicuous long!
term consequences[ This paper comments brie~y on
both issues[ Evidence is drawn mainly from studies of
herbaceous vegetation but there appear to be some
principles that apply more widely[

Dominants\ subordinates and transients

0887 British
Ecological Society\
Journal of Ecology\
75\ 891809

In order to estimate the consequences of a species loss

upon its host ecosystem\ it is necessary to know what
role "if any# the organism concerned plays within it
"Grime 0862^ Whittaker 0864^ McNaughton 0867^
Lawton 0883#[ In his pioneering attempts to de_ne
the functional roles of species within plant and animal
communities\ Whittaker "0854\ 0864# recognized that
a useful _rst step is to order component species
according to their relative abundance or productivity[
When large numbers of the resulting {dominance
diversity| pro_les are constructed it is possible to
begin the search for consistent associations between
the traits of species and their abundance in ecosystems
and communities[
For European herbaceous vegetation\ there is an
enormous fund of information on the abundance and
characteristics of component species in relatively
small "c[ 0 m1# vegetation samples[ Discussions of the
functional signi_cance of these data are available in
Grime "0862\ 0876#\ Grubb et al[ "0871# and Mitchley
+ Grubb "0875#[ Many ecological factors and plant
traits deserve consideration as potential determinants
of dominancediversity pro_les\ and it is clear that
controlling e}ects vary in detail from site to site[ How!
ever\ a generalization can be attempted and this is
summarized in Fig[ 0 in the form of an idealised domi!
nancediversity curve "sensu Whittaker 0854#[ This
suggests that within the majority of herbaceous veg!
etation samples three elements can be recognized\
each capable of varying in species richness and taxo!
nomic identity\ and here described as dominants\ sub!
ordinates and transients[ The dominants are usually
few in number\ taller and more expansive in mor!
phology and account for a high proportion of the
biomass "as seen for a grassland example in Fig[ 1#[
Many subordinates consistently co!occur with par!
ticular dominants and\ although they are usually more
numerous as individuals than the dominants\ they are

smaller in stature "Fig[ 1# and form a lower proportion

of the biomass[ In marked contrast to the subor!
dinates\ the transients are heterogeneous and lack
_delity of association with particular dominants[
They make a very small total contribution to the
vegetation and vary in number and in functional traits
to a great extent[ Most are represented only as seed!
lings or juveniles and a high proportion are species
that occur as dominants or subordinates in other eco!
logical situations "Table 0# often situated nearby[ In
passing\ it is interesting to note that formal procedures
used to collect and analyse data on the species com!
position of plant communities "e[g[ Clements 0894^
Braun!Blanquet 0821^ Bray + Curtis 0846^ Kent +
Coker 0881# often have the e}ect of under!recording
or discarding information on transients which\ from
a classi_catory viewpoint\ are frequently regarded as
{mis_ts|[ Table 0 shows that such excursions by species
appear commonplace when detailed and exhaustive
sampling procedures are applied] records from all
three of the sampled habitats include many species
that are more typically associated with other habitats[

Immediate effects of biodiversity

Can we generalize about the relative importance of
dominants\ subordinates and transients as deter!
minants of ecosystem properties such as productivity\
carbon sequestration\ water relations\ nutrient cycling
and storage\ litter quality and resistance and resilience
to perturbations< Both theory and experimental evi!
dence "Huston 0886^ Aarssen 0886# suggest that the
extent to which a plant species a}ects ecosystem func!
tions is likely to be closely predictable from its con!
tribution to the total plant biomass[ This {mass ratio|
hypothesis is implicit in many commentaries and
models relating to ecosystem function "Shugart 0873^
Pastor + Post 0875^ Huston + Smith 0876^ Grime
0876^ Sala et al[ 0885^ Huston 0886# and is dictated
by the laws of physics and chemistry that require that
large e}ects of autotrophs within ecosystems involve
major participation in syntheses\ and in inputs to
resource ~uxes and degradative processes[ It follows
that ecosystem properties should be determined to a
large extent by the characteristics of the dominants
and will be relatively insensitive to variation in species
richness in circumstances where this is attributable to
changes in the number of subordinates and transients[
It is important to specify that the mass ratio hypoth!
esis is restricted in application to the role of auto!
trophs in ecosystem processes[ When attention is
turned to other trophic elements\ such as parasites\
herbivores\ predators and symbionts\ the possibility
arises for ecosystem impacts that are less predictably
related to abundance[
A growing body of experimental evidence supports
the hypothesis that ecosystem properties are strongly
in~uenced by the characteristics of dominant plants[
In a comparative study of the resistance and resilience

893
Bene_ts of plant
diversity to
ecosystems

Fig[ 0 An idealised dominancediversity curve "sensu Whittaker 0854# for a small sample of herbaceous vegetation[ The
distinction between dominants and subordinates is based upon relative abundance and has been set at an arbitrary value
"09)#[ Transients are distinguished from subordinates by their failure to regenerate and persist in the vegetation under
scrutiny[

0887 British
Ecological Society\
Journal of Ecology\
75\ 891809

of _ve herbaceous communities to drought\ late frost

and _re "MacGillivray et al[ 0884#\ predictions using
traits measured in the laboratory were found to be
accurate when calculations were based upon means
weighted according to the relative abundance of each
plant species at each experimental site[ A similar con!
clusion was drawn from two recent investigations
"Wardle et al[ 0886^ Hooper + Vitousek 0886# in
which various ecosystem properties were found to be
strongly correlated with the functional characteristics
of the dominant contributors to the biomass[
There is also strong evidence that functional
di}erences between co!existing dominants can have
profound e}ects on ecosystems\ particularly in sus!
taining yield over periods in which there is ~uctuation
in climate or vegetation management[ From both
monitoring studies and experiments "Willis 0852^ Mel!
linger + McNaughton 0864^ Kemp + Williams 0879^
Grime et al[ 0874# there is abundant evidence that
di}erences occur between codominants\ in phenology\
photosynthetic mechanism\ rooting depth and repro!
ductive biology[ In some reported cases "Spedding +
Diekmahns 0861^ Armstrong + Eadie 0862^ Hooper
+ Vitousek 0886# such complementary exploitation
has been shown to confer a bene_t to productivity[
If the immediate in~uence of vegetation on the
properties of ecosystems is determined primarily by
traits of the dominants\ it is necessary to consider
what additional e}ects might be exerted by sub!
ordinates and transients[ First\ perhaps\ it should be
noted that there is no a priori reason to suspect that
such minor contributors must in~uence ecosystem

functioning^ their presence may simply re~ect the fact

that conditions prevailing in the past or present have
allowed them to be admitted[ However\ several sour!
ces of evidence suggest that some subordinate mem!
bers of plant communities "as distinct from transients#
ful_l roles that extend beyond that of mere adventives[
We may suspect that the consistent associations evi!
dent between certain dominants and their sub!
ordinates "e[g[ Fig[ 1# re~ect a complementary exploi!
tation of habitat resulting in a more complete capture
of resources and minor bene_ts to productivity[ In
some pairings of subordinates with dominants\ spatial
and temporal interlocking _nds precise morpho!
logical expression\ as in the case of the bryophyte
Brachythecium rutabulum which\ in winter\ colonizes
each fallen stem of the litter of the tall herb Urtica
dioica "Furness + Grime 0871#[ More often comp!
lementarity between dominants and subordinates
consists of circumstances in which the latter exploit
relatively unfavourable microhabitats[ These may be
expressed spatially\ as in the case of the creeping herbs
and bryophytes that occupy the shaded lower strata
of herbaceous canopies "Al!Mufti et al[ 0866^ Fig[ 1#\
or temporally\ as exempli_ed by the small winter
annuals and spring geophytes that make a minor con!
tribution to the biomass of species!rich calcareous
grasslands "Ratcli}e 0850^ Grime et al[ 0874#[
Further evidence of mechanisms of habitat exploi!
tation complementarity between dominant and sub!
ordinate members of a plant community is available
from an experiment "Campbell et al[ 0880# in which
the abundance of eight plant species in an exper!

894
J[P[ Grime

Fig[ 1 Vertical distribution of the leaf canopy of "a# four selected dominant and "b# four selected subordinate component
species in an ancient limestone pasture at Buxton\ North Derbyshire\ UK[ Canopy distribution in June was estimated by
measuring contacts with 264 randomly distributed\ vertical pins "S[ H[ Hillier\ unpublished data#[

Table 0 Ecological classi_cation of the species recorded in three distinctive and highly contrasted habitats sampled widely in
an area of 1399 km1 around She.eld\ UK[ All the species encountered within a particular habitat were classi_ed in terms of
their primary habitat "columns 25#[ Details of the sampling\ recording and habitat classi_cation procedures are provided in
Grime et al[ "0877#

0887 British
Ecological Society\
Journal of Ecology\
75\ 891809

Sampled habitat

Number of
m1 samples

Woodland
species

Grassland
species

Arable
species

Others

Total

Woodland on limestone
Meadows
Cereal arable

40
39
44

54
6
4

12
53
27

0
7
58

4
9
1

83
68
003

 Includes species primarily associated with wetland or skeletal habitats "cli}s\ walls and rock outcrops#[

895
Bene_ts of plant
diversity to
ecosystems

imental mixture was accurately predicted from inde!

pendent measurements of shoot and root foraging
by isolated plants growing in a standardized patchy
environment[ From this investigation it was con!
cluded that dominance was achieved by the devel!
opment of a coarse!grained architecture in which
main roots and shoots spread rapidly through a large
volume of habitat with rather imprecise concentration
in resource!rich sectors[ A complementary foraging
mechanism was recognized in subordinates^ here
resource capture was achieved by a precise but local
concentration of roots and shoots in resource!rich
patches\ a specialization likely to carry the penalty
of subordination and ultimately "in circumstances of
unrestricted growth and consolidation by the domi!
nants# risk of competitive exclusion[
We may conclude\ therefore\ that functional diver!
sity among dominants and perhaps also within sub!
ordinates is capable of immediate impacts on the
properties of ecosystems[ However\ it may be a mis!
take to narrow the search for bene_cial e}ects of
biodiversity to an examination of immediate e}ects^
the role of minor vegetation components may become
evident only when viewed in the long term[ The
remainder of this paper outlines two hypothetical
mechanisms whereby bene_ts of biodiversity may
accrue from intermittent e}ects of minor contributors
to the plant biomass[ The _rst hypothesis involves
subordinates\ the second transients[

Filter effects of biodiversity^ a role for sub!

ordinates<

0887 British
Ecological Society\
Journal of Ecology\
75\ 891809

So far in this paper it has been convenient to regard

the plant community as a stable hierarchy containing
dominants\ subordinates and transients[ In reality\ of
course\ communities experience ~uctuations in com!
position driven by seasonal and longer!term changes
in climate\ herbivory and vegetation management and
by the intrinsic dynamics of component plant popu!
lations[ Moreover\ from _eld observations and experi!
ments there is strong circumstantial evidence that the
persistence of subordinates in both grassland and
woodland vegetation is frequently dependent upon
periodic events "e[g[ droughts\ frosts\ ~oods\ wind!
throws\ grazing\ trampling\ burning\ coppicing# that
temporarily restrict the vigour and competitive e}ects
of dominant plants[ The literature contains many ref!
erences to circumstances where abatement of such
damaging events has led to expansions by dominants\
losses of subordinates and a rapid decline in species
richness "Tansley + Adamson 0814^ Thomas 0859^
Smith et al[ 0860#[ There can be little doubt\ therefore\
that plant species that habitually dominate particular
plant communities usually exert controlling e}ects on
the _tness of their subordinates[ However\ since it is
suspected that the immediate controls on ecosystem
properties are largely determined by the dominants it
is much more relevant to the purposes of this paper to

ask {Do subordinate members of plant communities

exercise controls on the identity\ functional diversity
and relative abundance of dominants<|
In order to review the opportunities for sub!
ordinates to control dominants it is necessary to con!
sider the long!term dynamics of vegetation and the
regenerative phases in the life cycles of dominants[
From studies of vegetation succession "e[g[ Watt 0814\
0836# it is established that continued dominance by
particular species is frequently determined by the suc!
cess of seedling or vegetative re!establishment fol!
lowing disturbance events causing mortalities of
dominants on either a local or catastrophic scale[ Here
it may be important to recognize that often the early
course of events following a disturbance is a tem!
porary expansion in the cover and vigour of sub!
ordinates[ This phenomenon is most obvious in forest
clearings where a dense low cover of shrubs\ herbs and
bryophytes may provide the context for regenerating
trees "Watt 0814^ Skutch 0818^ Marks 0863^ Bormann
+ Likens 0868#\ but similar phenomena have been
described for grasslands and heathlands "Oosting
0831^ Keever 0849^ Hillier 0889#[ Patterns of seedling
and vegetative establishment following disturbances
are not determined exclusively by regenerative traits
such as the size and number of propagules and their
dispersal\ dormancy\ morphology and physiology[
They arise also through complex interactions with
substratum conditions in which contributions to the
ground cover by subordinate plants may be expected
to have both positive and negative e}ects "Cavers
+ Harper 0856^ Ross + Harper 0861^ Grubb 0866^
Connell + Slatyer 0866^ Noble + Slatyer 0868^ Pickett
+ White 0874^ Bazzaz 0875^ Maguire + Forman 0872^
Burke + Grime 0885#[ Bene_ts to establishment have
been described in circumstances where seedlings sur!
vive in the shelter a}orded by low!growing shrubs\
herbs and bryophytes "Lawrence + Hulbert 0859^
Ward 0889^ Hillier 0889#[ Negative e}ects of shrub\
herbaceous and bryophyte cover on the establishment
of grassland and forest dominants have been observed
"Wardle 0848^ Niering + Goodwin 0851^ Webb et al[
0861^ Pons 0878# and it is widely accepted "Fenner
0881# that many small!seeded herbs\ trees and shrubs
are incapable of establishment where there is a closed
cover of vegetation[ There is some evidence that quite
inconspicuous subordinate members of the plant
community can exert a selective e}ect on seedling
populations of regenerating dominants[ For example\
in a microcosm experiment reported in Grime "0876#
an algal _lm on the soil surface suppressed the devel!
opment of small!seeded species but permitted estab!
lishment of larger!seeded grasses[ The signi_cance of
subordinates in plant communities therefore may
extend beyond any immediate contributions to the
carbon economy and nutrient dynamics[ Over the
longer term there appears to be a potential for sub!
ordinate members of a plant community to act as a
_lter selecting between di}erent potential dominants

896
J[P[ Grime

during the early phases of recolonization following a

disturbance event[ Such selection could operate on
the basis of variation in the seed reserves of dominants
and the associated di}erences in the capacity of their
seedlings to penetrate a low canopy "Grime + Je}rey
0854^ Westoby et al[ 0881#[ Alternatively\ according
to the characteristics of the ground cover the _lter
might discriminate between dominants that rely upon
rapid emergence and those which regenerate by per!
sistent juveniles "Marshall 0816^ Chippindale 0821^
Marks 0863#[ Controlling e}ects of subordinates
upon regenerating dominants may also occur through
more indirect mechanisms\ such as provision of sites
in which seed predation is reduced "Thompson 0876^
van Tooren 0877#[ In this review it has been con!
sidered prudent to restrict discussion of the possible
_ltering role of subordinates to rather direct e}ects
on the recruitment of dominants[ It would be possible\
however\ to include cases where the impact of sub!
ordinates arises from more complex phenomena such
as the maintenance of critical pests\ pathogens\ her!
bivores or mutualists "e[g[ Gilbert 0866^ Huston +
Gilbert 0885#[
Evidence of a _lter role for subordinates during
ecosystem re!assembly remains anecdotal[ There is an
urgent need for carefully designed\ long!term experi!
ments "e[g[ Ward 0889# to evaluate this phenomenon[

Founder effects of biodiversity^ a role for tran!

sients<

0887 British
Ecological Society\
Journal of Ecology\
75\ 891809

On _rst inspection\ transients appear to be irrelevant

to ecosystem function[ They occur as scattered indi!
viduals and many appear only brie~y as seedlings
that fail to survive[ Familiar examples in European
grasslands\ for example\ include annuals such as spec!
ies of Papaver and Poly`onum that occur as a legacy
of former arable cultivations or seedlings of wind!
blown or bird!dispersed herbs\ shrubs and trees[ Can
such minor and incongruous constituents of veg!
etation a}ect the functioning and viability of eco!
systems<
In order to explore the ecological signi_cance of
transients it is useful to identify the origin of these
individuals and to consider why some communities
contain a wider diversity than others[ This review
concentrates on the possible signi_cance of transients
as an indicator of the e}ectiveness with which poten!
tial dominants are dispersed across the landscape and
recruited into {suitable| ecosystems[ However\ it is
also worth noting that transients represent a neglected
subject in plant ecology and are worthy of studies
beyond the scope of this review[ In particular\ there
is a need to test the hypothesis that the accumulation
and persistence of transients in species!rich vegetation
is an indication that a low intensity of competition
prevails in such conditions[
The sources of the transients appear to be seed
banks in the soil and the seed rain from the sur!

rounding landscape[ This suggests that the transients

may provide useful information concerning the pool
of potential colonizing species at each site[ We may
predict that a diversity of transients signi_es the pres!
ence of a rich assortment of colonizers and a high
probability that\ in the event of habitat disturbance
or changes in management\ there will be a rapid
ingress of di}erent plant functional types\ some of
which may be capable of exploiting the new
conditions[ Here a speci_c example would be the ben!
e_t to woodland development where an abandoned
grassland already contains a diverse assortment of
tree seedlings[
E}orts to conserve biodiversity in Europe and in
many other parts of the world takes place in a frag!
menting landscape mosaic continuously disturbed by
natural events and by urbanization\ arable culti!
vation\ forestry and various forms of grassland man!
agement[ Successful conservation therefore depends
in part upon continuous movement of populations
and re!assembly of vegetation types and ecosystems[
The extent to which communities and ecosystems are
rapidly reconstituted is likely to be related to the res!
ervoir of colonizers\ many of which should be detect!
able prior to disturbance as transient constituents of
the existing vegetation[ Following Egler "0843# we
may suspect that the speed and completeness with
which ecosystem re!assembly occurs will depend upon
early colonization by appropriate dominants and sub!
ordinates^ late arrival may be expected to delay estab!
lishment of a species and may even exclude some
completely "Keever 0849^ Niering + Goodwin 0851^
Holt 0861^ Platt 0864#[ It follows\ therefore\ that the
decline in diversity occurring in many contemporary
landscapes is not simply a consequence of in situ losses
within communities[ Impoverishment may also occur
through a progressive failure in the processes of plant
dispersal and ecosystem re!assembly^ this failure
should be detectable as a decline through time in
the density and species richness of transients in plant
communities[

Implications for ecological theory

This paper has sought to connect recent studies of
plant diversity and ecosystem function "e[g[ Lawton
0883^ Naeem et al[ 0883^ Tilman + Downing 0883#
with another literature concerned with the mech!
anisms controlling diversity itself[ Reference to inves!
tigations such as those of Whittaker "0864#\
McNaughton "0867# and Grime "0862# reveals that
even in species!rich vegetation most of the plant
biomass may reside in a small number of dominant
species\ the characteristics of which are likely to over!
ride as ecosystem controllers the e}ects of more
numerous subordinate or transient components[ As
discussed by Huston "0886#\ this suggests a need for
cautious assessment of studies "Naeem et al[ 0883^
Tilman + Downing 0883^ Tilman et al[ 0885^ Tilman

897
Bene_ts of plant
diversity to
ecosystems

et al[ 0886# in which correlations are established

between species richness and ecosystem properties but
data are not presented with respect to the relative
abundance of component species[ Extreme care
appears necessary where di}erences in species richness
are not only correlated with di}erences in ecosystem
functions but are also confounded with contrasting
life histories and resource dynamics "e[g[ Tilman +
Downing 0883#[ However\ until the results of long!
term experiments measuring the functional charac!
teristics of natural and synthesized ecosystems of con!
trasted species richness and functional composition
are available\ the possibility of immediate e}ects of
species richness per se cannot be eliminated[
Even if the balance of evidence "Huston 0886^
Grime 0886# continues to shift towards the mass ratio
hypothesis and against the proposition that species
richness controls the immediate functioning of eco!
systems\ this does not mean that losses of plant diver!
sity should be viewed with equanimity[ Declining
diversity may be associated with less obvious impacts
that operate through failures in _lter and founder
e}ects[ In particular we suspect that there may be a
progressive loss of functions in circumstances where
vegetation patch dynamics and ecosystem re!
assembly continue against the background of a declin!
ing pool of colonizing propagules[ According to this
hypothesis the signi_cance of plant diversity in
relation to deterioration of ecosystem functions may
arise primarily from its e}ects on the recruitment of
dominants rather than any immediate e}ects of spec!
ies richness per se[

Acknowledgements
I am grateful to Sue Hillier\ John Hodgson and Ken
Thompson for permission to use data obtained in
collaborative projects\ and to Sarah Buckland\ And!
rew Askew and Suzanne Hubbard for assistance in
preparation of the manuscript[ It is a pleasure to
acknowledge the perceptive and constructive com!
ments of Sandra Lavorel and Michael Huston on an
earlier draft of this paper[ Some of the research drawn
upon in this paper was supported by the Natural
Environment Research Council[

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0887 British
Ecological Society\
Journal of Ecology\
75\ 891809

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Bene_ts of plant
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Received 06 November 0886
revision accepted 29 March 0887