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of the maximum-likelihood, neighbor-joining, and
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37 Gaut, B.S. and Lewis, P.O. (1995) Success of maximum likelihood
phylogeny inference in the four-taxon case, Mol.Biol. Euol. 12,
152-162
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Comput. Appl. Biosci. 8, 275-282

Biodiversityand the productivity

and stabilityof ecosystems
Kris H. Johnson, Kristiina A. Vogt, Heidi 1. Clark,
Oswald 1. Schmitz and Daniel 1. Vogt

R
esolution of the relation- Attempts to unveil the relationships The diversity-stability hypoth-
ships between the diver- between the taxonomic diversity, esis3 introduced the idea that
sity of life forms occupying productivity and stability of ecosystems increasing the number of trophi-
, ecosystems and the behav- continue to generate inconclusive, tally interacting species in an eco-
ior of those systems is a prime contradictory and controversial logical community should increase
directive of ecological research. conclusions. New insights from recent the collective ability of member
The problem ultimately encom- studies support the hypothesis that populations to maintain their
passes all questions about how species diversity enhances productivity abundances after disturbance. In
species coexist and how communi- and stability in some ecosystems, but not his presentation of the hypoth-
ties of populations influence eco- in others. Appreciation is growing for the esis, MacArthur3 implicitly recog-
system performance. Early theo- ways that particular ecosystem features, nized the transfer of energy from
retical discussions established the such as environmental variability and one trophic level to another as a
axiom that diverse, complex eco- nutrient stress, can influence biotic quintessential ecosystem function.
logical communities are the most interactlons. Alternatives to the Biomass accumulation is the intui-
stable’. Results of some field stud- diversity-stability hypothesis have been tively sensible, practical measure
ies in the 1960s and 1970s began to proposed, and experimental approaches of energy assimilation, and field
challenge the universality of this are starting to evolve to test these studies that followed used some
paradigm. Theoretical advances hypotheses and to elucidate the measure of biomass as the eco-
based on mathematical modellingl mechanisms underlying the functional system function response variable.
indicated that the nature of species role of species diversity. The formalization of the diversity-
interactions, rather than species stability hypothesis served as the
number alone, determines the original impetus for framing eco-
stability of ecological systems. Kris Johnson, Kristiina Vogt, Heidi Clark, logical questions in terms of the
Oswald Schmitz and Daniel Vogt are at the
Subsequently, several hypotheses relationship between diversity and
Yale School of Forestry and Environmental Studies,
about diversity and ecosystem stability. The hypothesis came to
205 Prospect Street, New Haven, CT 06511, USA
function relationships have been (kjohnson@minerva.cis.yale.edu). be popularly acclaimed as a law of
proposed. Recently, growing con- nature in spite of the fact that
cern over the loss of biodiversity results of some field studies failed
and new empirical evidence has to support it’.
prompted revisitation of the idea that species diversity The remaining hypotheses were introduced as alterna-
enhances the productivity and stability of ecosystems*. tives to the diversity-stability hypothesis. The rivet hypoth-
esis4 suggests that ecosystem resistance - the ability of a
Hypotheses and theoretical foundations system to absorb changes in abundances of some species
Four prevalent hypotheses (in addition to the null model) without drastically changing ecosystem performance (e.g.
are summarized in Box 1:the diversity-stability, rivet or rivet- biomass production)5 - can decline as species are deleted
popper, redundancy, and idiosyncratic response hypotheses. even if system performance appears outwardly unaffected,

372 0 1996, Elsevier Science Ltd PII: SO169-5347(96)10040-9 TREE uol. II, no. 9 September 1996
 REVIEWS

with potentially sudden and drastic consequences. Ehrlich

and Ehrlich4 presumed that it may not be known ahead of Box 1. Hypotheses for the functional role
time which, or how many, species deletions may cause eco- of species diversity in ecosystems
system collapse (Box 1). The redundancy hypothesis, as pre
sented by WalkerG, proposes that some species may be (4 (b)
expendable in terms of ecosystem maintenance if any
extant species can take the extinct species’ functional place
in the community. This idea originated with Ehrlich and
EhrlichJ as part of the rivet analogy; Lawton recently advo-
cated a useful distinction between these two hypotheses
(Box 1). The redundancy hypothesis can be viewed as a
redirection of the rivet hypothesis towards explicit consid-
eration of which ‘rivets’are likely to be functionally expend-
able within a given scenario of previous deletion@. Most
recently, Lawtonr introduced the idiosyncratic hypothesis,
which acknowledges that there may be no pattern or an
indeterminate relationship between species diversity and
ecosystem function. Results of experiments designed to Cd)
identify the functional significance of species diversity can
be interpreted to support one or more of these hypotheses.
The development of the alternative hypotheses traces
back to the diversity-stability hypothesis by association.
MacArthur3 was primarily concerned with the temporal con-
stancy of population numbers in the event of disturbance.
The contemporary definition of ecological stability refers to
the ability of a community to (1) resist change in its ability
to maintain an ecosystem function (resistance), and (2) re-
cover to normal levels of function after disturbance (resili-
ence)s. This refinement of the meaning of ecological stability
occurred partly through debate following MacArthur’s pro-
posall. Walkers referenced the historic mathematical chal- No. of species
lenges to MacArthur’s hypothesis’ in justifying his articu-
lation of the redundancy hypothesis. Lawtonr considered the A number of hypotheses about the functional role of species diversity within eco
logical communities have been proposed.
redundancy6, rivet4 and idiosyncratic7 hypotheses together
to address rates of ecosystem processes. McNaughton@ Diversity-stability hypothesls: this hypothesis [represented in Fig. (a)] predicts
and his colleagues extrapolated the diversity-stability hy- that ecological communities will increase in energetic efficiency (productivity), and
pothesis to the ecosystem level by treating primary pro- in ability to recover from disturbance, as the number of species in the system in
creases. According to this concept, deletion of any species from a food web will
duction rate as the response variable in species diversity
increase the susceptibility of the system to disruption via perturbation. MacArthur3
experiments. Tilman and Downing10explicitly state that the proposed this hypothesis in 1955 under the premise that energy flow in complex
redundancy hypothesis is an alternative to the diversity- food webs (meaning those featuring the greatest number of interspecific links or
stability hypothesis. Ah of the hypotheses in Box 1 are, there ‘connectance’) will be least disrupted by disturbance because alternative pathways
fore, phylogenetically related, and have come to be accepted for energy flow will be used. Elton’s restatement of this hypothesi+ is based on
the same principle and is also frequently cited as the origin of the idea. Insofar as
as contextually homologous by investigators of the func- the diversity-stability hypothesis has been accepted as an alternative to any of the
tional significance of biodiversity2,10-12.Indeed, ecosystem other hypotheses graphically portrayed here [Figs (b)-(d)]2.6.10-12, our graphical
resilience could very well serve as the ‘process’ for all of the caricature [Fig. (a)] is logically derived. In its original forms, the diversity-stability
graphs in Box 1. hypothesis did not include assertions of linearity in the effect of species richness
on ecosystem function.

Patterns in empirical evidence
Support for the idea that species diversity is crucial for
maintaining ecosystem functions has come primarily from
a microcosm experiment in a controlled laboratory facility
called the EcotrorPJ3, and from plant community responses
in field experiments conducted in herb-dominated systems
in seasonal environment&lOJ4J5. Examinations of other sys-
temsPJ7 have generated support for some of the alternative
hypotheses (Box l), suggesting that intrinsic characteris-
tics of particular ecosystems influence the functional roles
of their resident species and, therefore, of species diversity.
Among field experiments, findings about the functional role
of species diversity vary somewhat, depending on the
choice of state variables, the way that those variables are
measured, and functional distinction between the species
involvedQJ8.
Table 1 summarizes experimental approaches and the
results of some classical investigations into the influence of
Rivet hypothesis: introduced by Ehrlich and Ehrlich” in 1981, this hypothesis
[Fig. (b)] likens species in an ecosystem to rivets holding an airplane together -
the removal of rivets beyond some threshold number may cause the airplane, or
the ecosystem, to suddenly and catastrophically collapse. Explicit in their presen
tation is the appreciation that a few extinctions may go unnoticed in terms of sys-
tem performance because some species may be redundanP, generating a nonlinear
relationship between species richness and ecosystem function.
Redundancy hypothesis: advanced by Walkers in 1992, the redundancy hypoth-
esis [Fig. (c)] proposes that certain species have some ability to expand their ‘jobs’
in ecosystems in order to compensate for neighbor species that go extinct, sensu
Ehrlich and Ehrlich4. At the heart of this concept is the idea that species may be
segregated into functional groups; those species within the same functional group
are predicted to be more expendable in terms of ecosystem function relative to
one another than species without functional analogs.
Idiosyncratic hypothesis: Lawton’ included this model [Fig. (d)] to address the
possibility of a null or indeterminate relationship between species composition
and ecosystem function. This relationship would be expected, for example, in com-
munities featuring higher-order interactions.
All figures, except for part (a), are redrawn, with permission, from Ref. 7.

species diversity on the productivity and stability of eco-

systems, focusing mainly on terrestrial systems for brevity.

TREE vol. II, no. 9 September 1996 373

REVIEWS

In most cases, species diversity increased and rate of bi@ initial levels faster than in species-poor plots (Table 1) was
mass production decreased with successional time9JJJg-21. taken as support for the diversity-stability hypothesis. Since
In the first of two experiments by Tilman and colleagues in the theoretical mechanism behind the diversity-stability
Minnesota grasslandslo, nitrogen (N) fertilization ultimately hypothesis (Box 1) is that additional species have different
resulted in a decline in species richness, so the plots that niches and consequently contribute to productivity in some
accumulated the greatest biomass (standing crop) also con- additive fashion8J0, it is interesting that overall biomass de-
tained the fewest species. In a second experiment specifi- clined with increasing diversity in their studylo. Such dis-
cally designed to test for species diversity effects, Tilman crepancy between the mechanistic theory underlying the
et al. seeded plots with various numbers of grasses randomly diversity-stability hypothesis and the presumably support-
selected from a pool of native species, and observed a posi- ive empirical evidence testifies to the challenge posed by
tive diversity-productivity relationshipl5. In a novel experi- the pursuit of reliable knowledge of diversity and ecosystem
ment employing the Ecotron, Naeem et al. examined a suite function relationships. The disparity in the findings about
of ecosystem functions relative to differences in randomly diversity-productivity relationships between the Tilman
assigned species assemblages. One of their findings was and Downing study10 and the more recent experiments from
that primary productivity, measured as light absorbed by Minnesota’s and the Ecotronl3 (Table 1) has not been
plants and as carbon dioxide (CO,) fixation, increased with addressed.
plant species diversityl3. McNaughtong recently re-examined results of some early
Analyses of community stability also produced mixed studies of New York old fields that included observations on
results. The stability of plant productivity increased with spe- the responses of consumer communities, as well as those of
ties diversity in most studieslOJOJ1,as did retention of min- herbaceous plants, to pulsed nutrient enrichment (Table 1).
eral N in the most recent study from MinnesotaIs. However, Communities of arthropod herbivores and carnivores ap-
California annual grasslands displayed no effect of species peared, like plants, to increase in number and equitability
diversity on fluctuations in standing crop between yearsg, (evenness) of species with successional time. Herbivores,
while persistence of community types (measured as pro- unlike plants and carnivores, exhibited positive relation-
portions of grasses, forbs and other plants) in British hay ships between productivity and species diversity. In contrast
meadows declined with increasing diversity20Jl. to plants, the most diverse consumer communities were most
Tilman and Downing’s report of diversity-stability re- sensitive to, and slowest to recover from, the environmental
lations in Minnesota grasslands10 has received considerable changes caused by fertilization. This illustrates that the na-
attentionll,l*,22,23.This experiment was not originally de- ture of the perturbation and the selection of species for study
signed to examine the role of species diversity in maintain- can affect the outcome of diversity-stability experiments.
ing ecosystem function, and there was no control system for While the discrepancies apparent between theory and
comparison22. However, the observation that aboveground empirical evidence have led some toward skepticism of ar-
biomass in species-rich plots changed less and recovered to dently theoretical dialogues about the functional role of di-
versity (see discussions by
McNaughtonaand HairstorP),
it may be neglect of mecha-
Table 1. Summary of selected diversity/ecosystem function studiesa nistic thinking about species
Diversity/ Diversity/
interactions relative to eco-
Disturbance Diversity Productivity Stability productivity stability system function that fosters
Ecosystem type measure measure measure relationship relationship confusion. The basis upon
which MacArthur3 formed
California annual annual variation Sa sea SC(NPP)-1 plants (-) plants (0) the diversity-stability hypoth-
grassland9 (aboveground)
esis was implicitly a purely
New York annual variation; S/log, N; NPPa ANPP plants (-) plants (t) mechanistic blend of food
old field@ N-P-K fertilizer S and H’ (above- and herbivores (-) herbivores (-)
belowground) carnivores (-),(O) carnivores (-)
web ecology and ecosystem
ecology; as direct (consumer
Serengeti8 A precipitation; H’ NPP ASC NAa plants(t)
versus producer) links in a
grazing (aboveground)
community became more
Yellowstone drought; grazing H’ A relative NA plants (+)
numerous, interspecific com-
grasslands14 (abovziound) abundances
pensatory responses (a type
British fertilization; S A vegetation; plants (-) plants (-)20, of indirect interaction) be-
grasslands20Jl mowing; (abovziound) compositionzO (+)20.21
A precipitation ASC20,21
came more likely. Walker-6
considered such interpopu-
Minnesota droughVO; S SC ASC; plants (-)10, plants (+)I0
grasslandslo. AS5 (aboveground) recovery rateb; NAL5
lation compensation as the
(+)I5
NA= hallmark of redundancy.
soil fertilityc A soil fertility NA plants (t)
Frost et al.25 have begun to
Costa Rican annual S
tropical variation formalize this concept math-
forestlg ematically, explicitly identi-
EcotrorG S S NPP NA plants (t) NA fying negative interactions
related to resource competi-
aS, species richness: SC, standing crop; NPP, net primary productivity; N, total community biomass; A, change in parameter; tion as likely indicators of
H’, Shannon index; NA, not assessed; +, positive association; -, negative association: 0, no relationship. compensatory potential. In
bResistance was measured as ln@b/B&) and resilience was measured as In(b/B), where b is biomass at time t after dis-
turbance and B is biomass before disturbance.
the absence of such com-
CComponents of soil fertility included: available nitrogen (N), phosphorous (P) and potassium (K) content; and pH and cation pensatory mechanisms, one
exchange capacity. or another of the alternative
hypotheses (Box 1) is mecha-
nistically more plausible.

374 TREE vol. II, no. 9 September 1996


The question of how important species diversity is to eco-
system function boils down to consideration of how species
in that system interactQ5. The relatively consistent finding
from the studies of old fields and grasslands summarized in
Table 1 that stability increases with plant species diversity
reflects influences of seasonality on species interactions.
Herbaceous species in seasonal environments exhibit phe-
nology reflecting seasonality in availability of limiting re-
source+, especially water, N and phosphorous (P). Com-
petitive coexistence is achieved with temporal resource
partitioning through phenological differences that result in
a sort of relay from one species to another in dominance
of the vegetation throughout a season8J6. Periodic disturb
antes can also be important since they can ‘reset’ a seral
system before competitive dominants extinguish competitor
population@. Finally, in systems regulated by consumers,
the ability of herbivores to change their diets in response
to changes in relative plant species abundances and anti-
herbivore defenses also influences community dynamics27.
In this manner, we see that the intrinsic characteristics of
an ecosystem influence the feedback interactions among its
resident species. And, in the case of most seasonal herb
communities, a positive correlation between species diver-
sity and functional stability is predictable.
Observations about other ecosystems
Because the living and abiotic elements interacting to-
gether necessarily vary between ecosystems, the nature of
the interplay of those elements must also vary between sys-
tems28.The functions of species in ecosystems are therefore
expected to be system dependent. While this concept
seems self evident, and has long been acknowledged in dis-
cussions of other classes of ecological experiments29J0, it
has largely escaped mention in recent reports of diversity/
productivity/stability studies. This has resulted in liberal
extrapolation of results from some of the studies mentioned
above to the functional role of species diversity in generalrl.
However, research from other systems suggests that eco-
system functions may not be universally dependent on spe-
cies diversity.
For example, Cowling et al.16have contrasted the growth-
form redundancy of plant species occupying South Africa’s
fire-frequented Cape shrublands, called fynbos, with that of
adjacent forests. Fynbos vegetation is characteristically
flammable owing to soil infertilityl6, and the associated dis-
turbance regime profoundly affects the way that life forms
interact with one another and with the environment. Con-
versely, the adjacent forest ecosystems characteristically
have a longer disturbance frequency and tend to achieve a
greater degree of successional equilibrium. Cowling et al.16
classified plants into 11 functional groups based on growth
form, and compared the growth-form richness and species
richness for the two ecosystem types. These groupings can
be interpreted as surrogates for measures of trophic niche
width under the assumption that growth-form is related to
resource acquisition strategy.
The nutrient poverty of fynbos should set the stage for
intense exploitative competition for resources between co-
occurring species with overlapping resource needs. Selec-
tion would result either in niche partitioning and, ultimately,
adaptive radiation, or competitive exclusion, if the disturb-
ance frequency was long enough to allow communities to
approach equilibrium. This scenario would result in pheno-
typically distinctive, functionally unique species with either
exclusive fundamental or realized niches31. However, the
disturbance frequency mediates exploitative competition
between plants in fynbosl6, rendering some prominent plant
TREE vol. II, no. 9 September 1996
REVIEWS
species functionally redundant. Thus, Cowling et af.16argued
that functional redundancy apparently increased as disturb
antes became more frequent. Perturbation experiments for
explicitly testing the fynbos redundancy hypothesis with
regard to primary productivity and nutrient cycling have
not been performed, but are certainly feasible following the
approach of McNaughton*.
Environmental factors other than disturbance also can
influence the functional redundancy of different species. Vitt
and Cheer7 found that vascular flora do not drive ecosystem
processes in Canadian boreal peat bogs. Rather, vascular-
plant communities are downstream by-products of geo-
chemical processes largely independent of biotic activity
within the bogs. Plant communities assemble along gradi-
ents of pH (bryophytes) and organic macronutrients (vas-
cular plants). Nutrient supply depends on the chemistry of
the water flowing into the peat bogslr. Nitrogen and P have
been found to be particularly limiting to plant productivity
in these systems?.
Vitt and Cheeu classified bogs into three types: low, mod-
erate and high mineral (elemental cations) content. Plant
species richness was greatest for the moderate class bogs,
which were also the most temporally and spatially variable
with regard to water chemistry, especially N concentration.
Thus, plant species richness did not correspond to eco-
system dynamics in a manner supportive of the diversity-
stability hypothesis. An idiosyncratic or null relationship
between species diversity and ecosystem performance, es-
peciallywith regard to nutrient fluxes, better describes such
bog vascular-plant communities.
Are functional distinctions strongest at the species
level?
Central to questioning the role of species diversity in
maintaining ecosystem function is whether or not species
designations best distinguish functional groupings. The im-
portant problem lies in determining where in taxonomic hier-
archy functionally distinct phenotypes exist, Phenotypes
have a strong genetic determinant, and it is phenotype that
translates into functional role. Taxonomy itself has continu-
ally evolved to embrace new technology and conventions in
classifying biota. With recent advances in protein, chromo-
some and DNAisolation, genetics has become increasingly
important in species determination. However, explicit func-
tional distinction has not yet become an important taxo-
nomic criterion for species identificatior?. Some species
designations continue to be controversial and dynamic. For
this reason alone, species names seem unreliable as a basis
for differentiating functional groups.
Examples have been described where either genetic or
functional differences are prominent between taxonomic lev-
els other than species. Cossalter and Boyle33have found that
populations of the tropical legume Pterocarpus macrocarpus
differ from one another in resistance to drought stress. As
another example, limber pine (pinus flexifis) along the east-
ern foothills of the central Rocky Mountains exhibits a multi-
trunk growth form and, until recently, each cluster was con-
sidered a single, distinct individual. Carsey and Tomback34,
however, discovered that the trunk clump actually develops
from caches of variably related seeds constructed by the
bird, Clark’s nutcracker (Nucifragacofumbiana). In this case,
unusually high genetic variability occurs within and between
individual tree clumps. Additionally, Duarte et ~1.32recently
pointed out that plants of different ancestry above the spe-
cies level can be functionally alike. These examples from
plant ecology show that the genetically bounded phenotypic
distinctions that determine functional uniqueness may not
375
REVIEWS
conform simply to current systematic distinctions at the
species level.
MacGillivray and Grime5 are developing a systematic
approach for functional classification of plants. Their tech-
nique employs principle component analysis of morphologi-
cal traits in order to group species into functional types.
The traits reflect life history strategies related to stress tol-
erance and ability to compete for nutrients. To test the no-
tion that resistance and resilience of community produc-
tivity to various disturbances should be predictable based
on their classification criteria, they subjected communities
of grass species in various functional groups to an array of
realistic disturbances (drought, frost and fire). Results from
their field trials (not included in Table 1) generally con-
firmed predictions that resistance would increase and resili-
ence would decrease as community types changed from
ruderals to slow-growing, long-lived competitive dominants,
as expected during succession5.
Effects of phenotypic plasticity and mycorrhizae on
p/ant species redundancy
Phenotypic plasticity refers to the pronounced influence
of environment over expression of the genes that determine
growth-form or behavior. The presence of phenotypic plas-
ticity can blur functional distinctions between competing
species, especially those closely related, rendering them
functionally redundantsz. Plasticity, which appears to be
inversely related to genetic heterozygosityss, tends to be se-
lectively advantageous, and therefore most common, where
the environment varies unpredictably33. Thus, where com-
petition might result in competitive exclusion or adaptive
radiation into functionally unique life forms, phenotypic
plasticity provides a plausible mechanism for long-term per-
sistence of redundant species. This is a pattern that might
be expected between generalists with overlapping resource
requirements.
Mycorrhizal associations also muddy the functional dis-
tinctions between species because they buffer plants from
environmental stresses, especially nutrient deficiencies, that
might otherwise instigate intense competition. For instance,
different kinds of trees in mixed-species stands host many
of the same generalist species of mycorrhizal fungi, some of
which help each host tree to acquire N and P (Ref. 36). The
fungal symbioses ease interspecific competition by homog-
enizing the distribution of limiting nutrients, and by equal-
izing nutrient acquisition capabilities. Such effects certainly
increase functional redundancy (in terms of primary prod-
uctivity) of tree species, and appear strongest for systems
in nutrient poor, seasonably variable environmentsaQ7.
Coexistence of plant species under such conditions may be
attributable to processes other than divergence of trophic
niches. In such cases, species redundancy should be rela-
tively high, so tests of the species diversity-stability hy-
pothesis would probably produce different results to those
reported for Minnesota grasslandslo. Further study of the
mediation of influence of species diversity on ecosystem
attributes by mycorrhizae and other mechanisms begetting
functional plasticity can contribute immensely to empirical
resolution of patterns of diversity/productivity/stability
relationships.
Directions for the futwre
It is noteworthy, given the importance of belowground
processes to ecosystem function5v38- especially in grass-
land@- that roots have received little attention in the stud-
ies reported in Table 1. Only in New York old fields9 was
belowground production monitored. Focusing on below-
376
ground responses to perturbation can help ecologists detect
ecosystem-level changes because root morphology and bio-
mass are particularly sensitive indicators of environmental
stress38.
Furthermore, scientists focusing on interannual variation
in ecosystem attributes between experimental communities
under natural field conditions are resigned somewhat to de-
scriptive analyses because climatic effects are largely un-
controllable. Explicit tests of diversity-ecosystem stability
hypotheses, like all experiments, require that observations
about ecosystem function be made relative to variation in
independent variables, and that controls exist for those
independent variables. Interannual change in environment
and ecosystem attributes can occur for a system indepen-
dently of deviations from ground stateis. Assessment of de-
viation from ground state can be made only by comparing
observed responses in treatments with those in real con-
trol+*. The Ecotron offers unique potential to examine eco-
system attributes under carefully controlled conditions, and
the most recent experiment by Tilman et al.lsexemplifies a
controlled design for examining the effects of species diver-
sity manipulations under field conditions.
Finally, it is clear that identification of functionally dis-
tinct groups of organisms is necessary if ecologists are to
generate a priori predictions about ecosystem sensitivity to
extinctions5.6. For nutrient-related ecosystem attributes,
such as primary productivity, functional groupings will
reflect resource acquisition strategies of member species.
Studies of food webs have sometimes blackboxed species
into functional groups based on trophic level.941. However,
this approach may lead to underestimating functional dif-
ferences between species within a trophic level, as might
be seen with a keystone predator. In terms of ecosystem
functions involving the processing of nutrients, it is likely
that functionally unique species will be trophic specialists
and will exhibit relatively non-plastic phenotypes. Plas-
ticity, which should be an evolutionarily stable strategy
where resource abundances fluctuate unpredictably, can be
genetically programmed, or achieved through symbiotic
interactions between individuals of different species.
Functional redundancy between species is likely to be high-
est where plasticity or other mechanisms facilitate co
existence of species that otherwise compete for consum-
able resources.
Reliable knowledge about the functional role of species
diversity in ecosystems can only come from cause-effect
resolution through meaningful hypothesis-testing in con-
trolled experimentsi5JsJ4. The difficulty of designing and im-
plementing controlled experiments for hypothesis testing is
daunting. Most studies to date, with some exceptions8,13,15,19,
have generated only correlative conclusions about diversity-
ecosystem function relations because species diversity has
rarely been manipulated independently of ecosystem attri-
butes like succession and total nutrient abundance. The most
compelling evidence to date that species diversity can im-
prove productivity under certain conditions comes from
the innovatively designed experiments of Naeem et al. in the
Ecotroni3 and Tilman et al. in grasslandsis. These conclu-
sions will not apply to all ecosystemsl3. The challenge posed
by the task of identifying functionally distinct groups6 for
a given ecosystem is formidable in itself, but is also necess-
ary to undertake in order to generate meaningful predic-
tions about the effect of particular species deletions on
system performances. Experiments designed to determine
the types and strengths of interspecific interactions are sug-
gested to be the best path to take in determining which spe-
cies have the potential to compensate at the population
TREE vol. I I, no. 9 September I996

level for declines in or extinctions of other species for spe-
cific ecosystemsQ5.
The array of hypotheses in Box 1 illustrates a growing
appreciation that species diversity can influence the stabil-
ity and productivity of ecosystems in a variety of ways. Our
understanding of these relationships will continue to pro-
gress as ecologists identify the mechanisms determining
the degree to which taxa perform unique and critical func-
tions. Identifying the taxonomic level at which functional
distinctions might exist for a given ecosystem remains a key
consideration in this pursuit. The approach to functional
ordination based on morphological traits currently being
developed by MacCillivray and Grime5 offers promise as a
means of predicting the functional importance of diversity
within the context of a specific ecosystem.
Acknowledgements
This manuscript was greatly improved over earlier
drafts by following suggestions made by John Lawton,
Maria Uriarte, Andrew Beckerman and an anonymous
reviewer. This review was supported in part by National
Science Foundation grants to Yale University, USA under
the Terrestrial Ecology Program with the University of
Puerto Rico and the International Institute of Tropical
Forestry, US Forest Service, as part of the Luquillo
Experimental Forest Long-Term Ecological Research
funding to K.A.V. and D.J.V., and by grant
NSF-DEB-9508604 to O.J.S.
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