A New Model for the Continuum Concept Author(s): M. P. Austin and T. M. Smith Source: Vegetatio, Vol. 83, No.

1/2, Progress in Theoretical Vegetation Science (Oct. 1, 1989), pp. 35-47 Published by: Springer Stable URL: http://www.jstor.org/stable/20038482 Accessed: 18/05/2010 23:00
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1989. 83: 35-47, Vegetatio Publishers. 1989 Kluwer Academic ?

Printed

in Belgium.

35

A new model

for the continuum

concept

M. P. Austin1 & T. M. Smith2 1 Division of Wildlife & Ecology, CSIRO, Group, Research 2Ecosystems Dynamics Canberra,
Accepted

P.O. School

Box

2602 Australia; Canberra, A.C.T. 84, Lyneham, Sciences, Australian National University, of Biological

A.C.T2600
12.04.89

Australia

Keywords:
curve

Environmental

gradient,

Fundamental

niche, Niche

theory, Realized

niche,

Species

response

Abstract of the is presented after considering the implications concept is a spatial concept dependent and current niche theory. Community controversy community/continuum on landscape is an environmental the continuum space. concept pattern while referring to an abstract are ill-defined and the assumption of competition When applying niche theory to plants, the mechanisms A reformulation of the continuum curves for species unrealistic. of bell-shaped response on the pattern of response testable propositions Eight presented gradients. response 1. Environmental 2. The curves. are to environmental of vegetation gradients or b) direct physiological are of two types, a) resource gradients gradients is a series of similar nested niche response of species to resource gradients fundamental is a series of separate, niche response of species to direct gradients 3. The fundamental

curves are such that they curves. 4. Species fundamental response response overlapping independent, in some part of the environmental have a relative performance space. 5. The shape of the advantage even bimodal but predictable from the fundamental realized niche is variable response given the other to the response 6-8 describe shapes of emergent community properties species present. Propositions is bimodal, dominance trimodal and standing crop unimodal. environmental gradient; species richness Detailed of these propositions comparisons and Tilman. These and Whittaker, Grime, of plants and vegetation. properties are made theories with are the alternative incomplete theories several of Ellenberg, generally Gauch accepted lacking

Introduction science can be defined as the study of Vegetation those processes which determine thepatterns of com position and emergent properties (e.g. species rich ness) observed as 4a coherent as principles in vegetation. Theory can be defined used group of general propositions of explanation for a class There of is (Macquarie Dictionary).

with vegetation little theory associated relatively nor have vegetation science given this definition, to develop scientists actively sought theory. The on Theory of and Models symposium Uppsala Vegetation Science' (Prentice & van der Maarel

1987)marked amajor shift and recognition of this

one of us (Austin At that meeting, deficiency. tested a series of propositions 1987) arising from forward by Gauch & the continuum theory put

phenomena'

36 Whittaker These were propositions in eucalypt forest in The evidence for the continuum

(1972).

rejected for canopy south-eastern New more

postulating theoretical in a more

positive new hypotheses or new, more robust In this paper, we attempt frameworks. to re-express the continuum theory of vegetation robust detected

species South Wales, Australia. The of scientific research is aspect

Cottam

presented by is less than con (1966) as was out by Daubenmire vincing pointed can be resolved without The controversy (1966). if the two ideas of a con repeating old arguments & Mclntosh tinuum are based on different and a community frames of reference. Fig. 1 shows a incompatible

patterns vides mechanistic nomenological

form, which a) allows for the in the eucalypt forests, b) pro rather than phe processes to account testable for the hypotheses the theoretical pro theory the for should

in an area with 4 species up a mountain associations which be present. Species might to their frequency due the recognised along transect transect BC are A, AB, and CD B, C, D with the combinations These being regarded as ecotones. of co-existing composed species

descriptions

and, patterns c) presents can be deduced which from advanced. positions A re-formulation resolve a) lingering problems

'communities'

of continuum from

are a consequence of the spatial pattern of the If we examine the altitudinal landscape. gradient however we find a continuum each other replacing with increasing through an adjacent were 30 m of species regularly in the sequence A, B, C, D altitude. If the transect were taken area where the first bench

controversy, community between sharp boundaries

example,

continuum/ the types

vegetation

often observed in the field (Austin 1985,1986),

in current continuum and apparent b) problems to vegetation, niche concepts e.g. applicable to symmetric response the skewed as opposed curves of species 1986, 1987, and in (Austin press), between c) differences of other theories (Austin We 1986), continuum vegetation theory and

AB would lower, then the combination rare. At 170 m only species become immediately A would be present. Similarly if the second bench or tableland BC would were become 30 m higher the combination common. On such a transect be A, B, BC, Communities

the 'communities' C and D with or associations

recognised would ecotones AB and CD. are

for example issues

organization those of Tilman in the following

(1988) and Grime (1979).

treat each of these sections:

a function of the landscape examined. Abrupt changes or gradual transitions on the landscape may occur depending pattern. groups of species can be recognised Co-occurring for any particular region with a recurrent pattern of landscape. would Many phytosociologists such associations recognise tinuum (Ellenberg 1988). munities is useful for as noda within these Labelling communication a con com and

Continuum/Community

controversy

no major debates are now conducted in Although the ecological literature regarding whether vege or in terms of continua tation can be described (Austin (ex 1985), other ecologists still modellers and theoreticians) perimentalists, etc. on in designing models experiments, persist communities the assumption that communities while exist and can be (Austin in press), recognised still define associations gists Maarel phytosociolo van der (Westhoff&

of these communities research, but extrapolation accurate only if the regions to other regions will be of environment have similar landscape patterns and climate. The continuum in relation to altitude will in another between con region growth tinue to exist and be applicable the correlations provided

vast majority of applied 1978). The of nature concerned with management ecologists reserves continue to use the term community.

variables (e.g. rainfall and tempera influencing and altitude do not change (Austin 1980). ture), from this that: We conclude 1. The continuum concept space, environmental applies to the abstract to any not necessarily

37

600

400 Altitude

(m)
200

ENVIRONMENTAL DISTRIBUTION OF SPECIES

SPATIAL DISTRIBUTIONOF SPECIES

Fig. continuous 1. Patterns of co-occurrence variation depends on a landscape an indirect environmental along gradient species A plant is a landscape and altitude of composition concept community along gradient. on the frequency in a particular of environmental combinations landscape. of four altitude, recognition note of

communities

distance geographical indirect environmental 2. The concept species can

on the ground

or to any

version

landscape environmental

gradient. of co-occurring of a community to a particular only be relevant of and its pattern of combinations variables ;community is a land

(1972) where 'major' species are regularly the gradient with optima evenly distributed along 'minor' species are independently spaced while each stratum e.g. trees, shrubs and herbs of composition with either regu distributions and with the species or strata either independent evi there is insufficient these possibilities,

distributed, c) stratal continuum (Austin 1985) in

which has a continuum

scape property. For a theory of vegetation to be generally appli cable itmust be expressed in terms of an environ mental tion. environmental space defined by causal variables whose of loca values are independent The

lar or irregular between relationships correlated. dence

Currently, to distinguish between

although Austin (1987) has demonstrated for one

type that canopy species optima are vegetation a temperature not regularly distributed along gradient. There views mal niche is a close correspondence of the vegetation continuum consider species ecologists between and how in relation these ani to

of which three continuum concept, variants can now be recognised (Austin in press) fulfils this criterion but one of the major defi of this theory has been the failure to ciencies adequately define an environmental gradient.

Current Two

Continuum reviews

theory of continuum the point of the theory (Austin that there are three basic and concept: continuum their limits

(1984) 'Indi theory. To quote from Giller resources of one species using marginal as cannot exploit them as efficiently presumably viduals individuals resources model of other species for which these are nearly is summarised (our italics). This optimal' in Fig. 2a, and can be re as the realized niche of species b being region of its fundamental from species a and c. This for Gauch & the reason

recent

1985 in press) make variants possible a) Gleason's with species independently

original abundance distributed

individualist optima

expressed restricted niche

to the optimal

along an environmental

gradient (Gleason 1926), b) Gauch &Whittaker's

by competition is probably concept

38 Whittaker have that canopy (1972) suggesting species modes regular (optima) along environmental contrast In (Mueller gradients. Ellenberg realized curves niche common in theoretical theories.

pseudo-gaussian treatments of There niche are are and

and

continuum

Dombois & Ellenberg 1974) proposed that the

niche (ecological in his terms) response take any one of a number of shapes (e.g. the niche theory's pro Fig. 2b and c) including for the posed response (Fig. 2a). His explanation could occurrence that the of skewed and bimodal effect of competitive could restrict responses was better adapted

mechanistic

to current three major weaknesses are non continuum theories: l)they and the mechanisms of competition 2) the curves bell-shaped pseudo for both fundamental

undefined;

a species to those environ species at the extremes of its tolerance provided ments it had an advantage in that environment. Most appear (Austin 1987 in press) species responses to be skewed, contrary to the bell-shaped

are unrealistic for responses resource the environmental plants; 3) and/or are ill-defined to all and the response gradients is assumed in shape. identical gradients a new model We of the continuum propose of vegetation which concept composition takes account of these problems. citly expli

response gaussian and realized niche

New

model

for a Continuum

Environmental Three

gradients of environmental gradient can be

types

recognised (Austin 1980):

1. Indirect complex altitude. environmental gradients The influence gradients of Whittaker of are the factor (1978) such as an altitudinal

is through variables like temperature gradient and rainfall which have a direct effect on plant and have growth complex location-specific correlations with altitude. Relationships based on such gradients cannot be extrapolated beyond measured 2. Resource the environment where and are not considered they were further.

^~\

RESOURCE GRADIENT

for plants are those where gradients the varying resource is consumed by plants in order to grow. There are only a limited number of such resources for autotrophic organisms: light, water, carbon dioxide, oxygen and essen from tial mineral nutrients. These differ

COMPETITOR REALIZED FUNDAMENTAL

resource size

gradients in that plants niche

for animals show

such

as food

TYPES OF SPECIES RESPONSE CURVES

an environ curve along of species response Fig. 2. Types niche realized mental (a) classic niche concept with gradient of under optimum conditions, (b) and (c) Ellenberg's concept extreme of fundamental niche conditions under realized

inclusive The

response

of a limiting that due to toxicity at levels beyond under natural conditions normally experienced decline

responses of plants is generally taken to be response type with an eventual

or less only more to these resources.

niche (b)with bimodal variable shape (c) restriction to one

extreme only.

(Fig. 3a).
3. Direct gradients for plants are those that have

39 in the absence of competition,

plants

herbivory

and disease (Gauch&Whittaker 1972; Whittaker

(/) c o o. a)

1978). Ellenberg suggested that plant species have similar optima and physiological response curves on the basis of his early experiments (Ellenberg

1953,1954; Mueller-Dombois & Ellenberg 1974).

(or inclusive), highly-productive curves (a in Fig. 3a) having species response which include or at least have the same range as Resource gradient those mental gradients However of less productive species (b, c in Fig. 3a). we also conclude that species funda responses are neither to direct inclusive environmental or nested, species and different optima Species nested fundamental niches with will often tend to be

? u o o

a a) *u a) a

(/)

separate showing distinct individualist ranges of response (Fig. 3b). This evidence leads us to put forward two fur ther propositions: niche response Proposition (2) The fundamental of species to resource gradients will take the form of a series of nested curves. response Direct
Fig. 3. sponse

environmental

gradient
re

Proposition of species

(3) The fundamental to direct environmental

niche

response gradients will independent,

differences Proposed to (a) resource and

in species fundamental (b) direct gradients.

take the form of a series of separate, curves. response overlapping The evidence based on biomass

physiological impact on plant growth but are not consumed. The two clearest exam are air temperature and soil pH which ples rates and the maintenance of govern growth

a direct

for these propositions is generally curves, i.e. based on response growth rather than some integrated fit vegetative ness measure. The propositions may not therefore apply to fitness measures success and reproductive involving (Bazzaz germination pers. comm.).

plant physiological integrity.The fundamental

response different optima The first may being show differential regimes along temperature spaced to adaptation with species the gradient of

Smith & Huston

model

(1989) have developed a

(Fig. 3b).
continuum of our proposition theory is therefore: re-expression

of plant dynamics which examines patterns of plant functional to environmental fac response tors (and associated at life-history characteristics) the level of the individual plant. They define a 2-dimensional environ simple (light* water)

to Proposition (1) Environmental gradients a continuum can be applied are of which theory two types a) resource gradients, b) direct physio logical gradients.

mental

of growth space and examine patterns to the availability of these two resources. response The approach is based on cost-benefit analysis on two specific and focuses trade-offs: (l)the trade-off between high and resource conditions rate under growth or light) water (either to continue and photosynthesis conditions 1968; Grime of low 1977; maximum

Plant physiological There has been about

response little discussion the in the continuum response of

the ability growth (i. e., survive) under resource availability (Parsons

literature

fundamental

Chapin 1980; Orians & Solbrig 1977; Bazzaz

40

1979; Huston & Smith 1987; Turnan 1988), and

(2) the limit to the combined ground tolerance a plant can

unlimited tional

and we combinations

need

of the func knowledge of viable (cf. properties

exhibit to low availability of both light (above

and water resource) (below-ground see Fig. 4. These patterns of trade-off resource), are a function of an array of constraints ranging from biochemical of the whole lar discussion to carbon allocation at the level

Tilman 1988).

Plant

realized

niche (Ecological

response)

1988 for simi plant (also see Tilman for light*nitrogen). The implication is that a series of physiological/ of these patterns morphological adaptations in different formance optima resource tion: fundamental (4) Species Proposition curves will be such that in a particular the resource space response portion of space. This leads define relative positions to a fourth proposi per in the

Austin (1987), in testing a series of propositions

regarding the species response curves put forward

by Gauch & Whittaker (1972), showed that the

skewed with response curves were predominantly for the tails towards more mesic conditions on a temperature in gradient eucalypt species A review of studies with south-eastern Australia.

oldfield species (Tilman 1987) and aquatic species (Wilson& Keddy 1985) shows a similar predomi
nance of skewed It is necessary the fundamental become under Austin strated species culture the (Austin in press). to provide mechanisms whereby curves

performance for trade-offs need to be The exact possibilities examined carefully. Some species of plants appear to low moisture and able to adapt simultaneously low nutrient conditions. Grime (1979) has drawn com to these 'stress-tolerators' which attention bine slow growth The a variety of distinctive and life-history physiological and of physiological combination rates with properties of plants are not

a species will have a relative over other species. advantage

curve is modified to response curve observed realized response of competition, and conditions herbivory

disease. Experiments

(Austin 1982; Austin &

morphological, properties. morphological

etal. 1980; Austin 1985) have demon in polyculture, that performance (multi can be predicted from mono mixture)

(Fig. 5). In a trade-off situa performance in decrease tion similar to Fig. 4 the progressive for an extreme-adapted advantage physiological species ductive response until it is outperformed give species would curve for the by the more pro rise to a skewed niche. The and con the extreme productive

Biomass

realized

response would be steep towards have a long tail towards the more ditions. ecological \ Resource Measures of relative

prediction studied (Austin the suite of species sequent tests of these relationships erties that both morphological of the species

and physiological are necessary to make the performance on and the results are totally dependent 1982). Sub have shown

and physiological prop are needed to predict the competition (Austin

outcome
Resource A in funda trade-offs types of species Fig. 4. Three probable Growth and sur mental response. (a) Species physiological vival under low levels of resource A, but not B. (b) Growth of resource survival under low levels and B, but not A. under (c) Maximum growth to survive under low bility high levels of resources resource level A or B. but ina

of multispecies

etal. 1985; see also Gaudet & Keddy

can be developed further proposition observations and experiments:

1988). A
from these

Proposition

(5) The

realized

(ecological)

re

can mixtures sponse of a species in multispecies from its fundamental be determined (physiologi

41
_ ? GRASSES

that relative growth physiological

rate is the predictor of relative not biomass. performance

Emergent
0:5 LO

(community) emergent

properties in

Several

, r- EUCALYPTS

the vegetation number of species co-existing (species richness), the degree of dominance exerted by an individual the total biomass (standing crop) the vegetation. There is an expand by produced ing literature on the causes of species diversity species, and

can be observed properties a number of species: comprising

(e.g.Grime 1973;Huston 1979; Brown 1981), but

Relative Fig. 5. Examples of physiological the prediction of performance.

realized (eco species in multispecies mixture from the species response logical) in monoculture. fundamental response (physiological)

of general application remain precise mechanisms elusive. A recent empirical study (Margules et al. 1987) has shown that complex patterns of species to environmental in relation richness gradients several emerge when are considered variables different environmental Austin simultaneously. to a number of patterns that

cal) response

plant growth mation is available munity. The

of whole (i.e. an integrated measure inmonoculture) provided such infor for all species in the com

(1980) drew attention these community show along environ properties in total biomass mental and domi e.g. gradients nance. Gauch & Whittaker were unable to (1972) any pattern of recognise environmental gradients, who demonstrated (1973) between presents richness of these about species richness contrary a strong along to Grime

of this proposition is that the corollary shape of the realized response curve is a function of the other species present and may easily show forms. very complex The mechanisms associated with this predic tion are presumably of competitive exploitation the resource at low levels, the interaction between resource exploitation and light com competitive

relationship species richness and pH. Grime (1979) a unimodal between relationship species and standing biomass plus litter. In view

levels petition with canopy closure at intermediate and at high, toxic levels interaction between com for light and physiological of tolerance petition Direct 1982; Tilman (Austin 1988). of will show greater partitioning gradients the gradient due to differential responses along with the relative per physiological adaptations formance growth Grace determined relative being by plant rates and ability to form a closed canopy. shown that relative (1988) has recently performance at an early in monoculture of growth is a stage the realized toxic conditions

results, any propositions conflicting are necessarily these community properties without further research. speculative

richness will show two (6) Species Proposition on both at maxima intermediate positions resource and direct gradients, these will be at positions (e.g. of either deficiency and the most gradient) This intermediate extreme values between or toxicity on a resource

physiological

equable value. curve of to the bimodal is equivalent in Fig. 6a. It is probable richness that species curves for functional groups (e.g. richness species ore tress) or for annuals, ephemerals, epiphytes taxonomic groups (e.g. Eucalyptus, dipterocarps, or mosses) will show the clearest pat terns. Grubb additional informa (1987) provides tion and discussion that different life suggesting

(Austin 1982) better predictor of in mixture formance logical performance

per ecological at a later stage than physio at that time. This suggests

conifers

42 COMMUNITY PROPERTIES SPECIES RICHNESS the opposing extremes of deficiency and toxicity.

Proposition (8) Vegetation standing crop (total

curve to shows a bell-shaped response biomass) environmental gradients (Fig. 6c). This may appear self-evident but the implica tions need to be explored. If total productivity is then it appears unlikely that a highly constrained, can support numer low productivity environment ous different growth forms or survival strategies.

% DOMINANCE

The maintenance resource trees

cost

and

absolute

amount

of e.g. low

required to support large life-forms in their occurrence may preclude environments.

resource These

concern plant-environment propositions and they are essentially static con relationships disease and dis cepts. The roles of herbivores, turbance are not addressed, neither is plant repro or senescence, and yet the duction, persistence TOTAL BIOMASS levels of these be factors influenced by It has been hypothe gradients. to herbi sized that the evolution of plant response has been directly influenced vory by nutrient environmental themselves and plant variables or even determined can

availability. Grime
have suggested advantageous However, of plants
along

(1977) and Chapin

low nutrient

(1980)

that low maximum in

growth rate is environments. the ability

GRADIENT ENVIRONMENTAL
Fig. 6. Postulated an environmental patterns gradient. for community properties

Thus, following herbivory. rates have slow growth inherently plants with that reduce herbivory evolved chemical defenses (Bryant et al. 1983) suggesting a gradient of herbi vory parallelling that of nutrient availability.

low growth to regrow

rate also reduces

forms will ness along

show different environmental

patterns

of species

rich Alternative There theories of con

gradients. the maximum pro Proposition (7) Dominance, by a single portion of total biomass represented under extreme species will show three maxima, conditions may exist where (or very few) species only where and under optimal conditions com determine processes species follow suggestions of one

community

has been

little active

development

tinuum theory since the 1960's (Goodall 1963; Whittaker 1967;Mclntosh 1967)with the excep
tion of the statement by Gauch & Whittaker have however (1972). Independent developments of occurred which actively concern the behaviour We consider here four along gradients. vegetation specific sets of theoretical concepts, species those of

competitive These

position (Fig. 6b).

propositions

Grime (1973, 1979; see also Tilman 1982) with

resource or direct, that any gradient, recognition has two extremes of stress which can not be col into a single stress value as the processes different at 'stress' are fundamentally determining lapsed

Ellenberg

(1953,

1954; Mueller-Dombois
regarding (1979) concerning species

&
pri

Ellenberg 1974) curves, of Grime

response

43 in vegetation, of for survival mary strategies nutrient-ratios Tilman 1982) (Tilman regarding

and light/nitrogen gradients (Tilman 1988) and of

Bazzaz (pers. theories/hypotheses put forward positions The earlier Whittaker) environmental make no We comm.). in relation here. Gauch and (Ellenberg, distinction between special Whittaker though the indirect nature of his (Table on and 1). Grime two of types examine their to the eight pro a E

workers

gradients, recognised gradients concentrate

(1978) clearly factor-complex Tilman both gradients,

and productivity. They and direct the resource collapse Grime into a single dimension. spaces gradient stress and with low production (1979) equates con hence with stress tolerator species. Tilman disturbance effectively determine will nutrient availability (resource) a productivity and that light gradient (at ground level) as a resource will be negatively correlated with the productivity gradient. siders that There assume carbon same does not appear to be any reason to all resources, e.g. light, water, oxygen, dioxide have the and mineral nutrients, type of productivity gradient. This needs The failure to distinguish direct physio dif fundamentally must based

Phosphorus Fig. 7. The different direct environmental shaped evolved. environmental possible gradients spaces combinations will determine which of resource and

within

the differently are species

with

more-or-less

gradient will give a as two the space rectangular will be largely independent and all pos gradients sible combinations would be viewed as likely to a direct temperature occur resource (Fig. 7a). The combination of and the two gradients ents a more complex picture a supra-abundance of water, tation can not such as water light pres (Fig. 7b). If there is then terrestrial vege

research.

as having logical gradients ferent properties e.g. they are not consumed, lead to limited generality of any predictions on a purely possibilities environmental combination
1. Nature

production-gradient are numerous when space is considered of a resource gradient

of environmental

The concept. a 2-dimensional (Fig. 7). The in phosphorus

different survive; fundamentally are and morphological physiological adaptations The combination of low light and low necessary. moisture is extremely rare and imposes difficulties for terrestrial plants. Adaptive trade-offs have not been achieved. of a direct gradient such as phosphorus laws. Available soil phos The possible combinations such as pH with a resource

Table

gradients. distinction distinction as to type as to type

Ellenberg Gauch Grime & Whittaker

No No

a) Stress/Productivity
b) Disturbance

by chemical phorus is controlled by the soil pH and is increas at low and high pH (Russell ingly less available The range of feasible combinations of other 1973). resources needs to be carefully examined, because there are biological to many variables. constraints Plant ot plants' response to herbivores response availability. Plants environ in

is governed

Tilman

a) Productivity
b) Disturbance a) Resource

Bazzaz

b) Disturbance Austin & Smith

a) Resource

appears related to nutrient low nutrient, hence low ments while plant

b) Direct

productivity to avoid grazing, may have adaptations in productive, environments high nutrient to herbivory may include high

response

44 rates. Fire regrowth both environmental is a function of theoretical models is ameasure of fitness. Current

and vegetation There are therefore limited combi productivity. nations of fire and other environmental variables but with in addition there the dependent The fundamental considered relationship plant biomass. of species has not response very extensively by theoreticians variable resource levels. This is a feedback

frequency conditions

(Table 2) are speculative. propositions Response as distinct to direct from resource gradients has barely been considered gradients (Table 2). The size of the volume or range of fundamental response within a resource space has similarly

been

been ignored (Table 2) though this will affect the range of applicability of Tilman's models (1988).
Ecological and there tory data responses are only now being

(Table 2). Tilman is very explicit but restricts his

consideration to non-toxic for many natural situations but may be reasonable the frequent occurrence of water-logging in many moisture habitats would need to be taken high as would into account, in high photo-inhibition light environments. ments at less than excessive occur (Bradshaw etal. 1964). The such direct declines models. should The be theoretical Declines in growth in experi levels also nutrient of

described adequately for species (Austin 1987)

is an urgent need for further confirma studies. Few if any studies have adequate to rigorously test alternative hypotheses

(Table 3). Tilman

example great from his

(1987) provides one clear

survey skewed a of old fields, where curves are response

implication into incorporated volume of the resource/

of variety apparent, contrary to his theoretical assumptions. on patterns of There is no common agreement richness species along gradients (Table 3). Grubb

space gradient allows it to function growth,

that a species' physiology in is usually unknown for yet what is required for

vegetative

(1987) summarises many of the alternative possi ble causes of species richness including age, area, and disturbance, factors isolation, glaciation

Table

2.

Patterns

of fundamental

response: fundamental to a resource

alternative

propositions. fundamental to a direct gradient of fundamental in resource

Species response gradient Similar Variable Inclusive?

Species response

Ranges response space

environment

Ellenberg

optima shape

As

for resource

gradient

Gauch & Whittaker

Grime

Not considered
Not considered

Not

considered

Not

considered

Not

considered

Not

considered

Tilman

Law Nested

of diminishing

returns,

Not

considered

Not specified beyond threshold level

for growth

Bazzaz

Variable Nested

shape,

Various

Limited

range

Austin & Smith

Similar

Optima,

Dissimilar Variable Independent

optima, shape,

Limited range
Certain combinations non-viable of

returns Law of Diminishing at high levels, with Decline Nested

responses

45

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G S

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46 which attention certain which most are of gradients. of species He draws of variables. situation in an experimental from physiological

independent to observations

richness

Species can be

response predicted

life forms appear productive

and forest (climbers to have maximum richness environments. Life

herbs) in the and

response (Fig. 5; Austin 1982; Grace 1988), but

the context-sensitivity of the prediction, given to be found further simplifying need concepts before prediction of the composition of natural we dis will be possible. Although to two types of the fundamental response tinguish in differences gradients, we cannot yet distinguish to the two types of the ecological response vegetation gradients. The challenge devise will discriminate outlined for vegetation is to scientists tests and experiments which observational between the various authors. The theoretical of different set of pro here replace those of Gauch & they are a mixture of mechanis plant physiological

forms

particular C4 plants)

categories physiological can be expected to

of plants (e.g. show complex more et al. than 1987;

response patterns particularly one gradient is considered (Margules is substantial Minchin 1989). There about extreme the occurrence of more environments, a dominance high recent

when

agreement in dominance authors also

in productive maximum suggest further examination, This needs environments. can this suggestion when be maintained tropical is so species rich? Similar differences rainforest apply The to single species of vegetation dominance and total with

propositions positions Whittaker tic

biomass (Table 3).

concept as a continuum and structure continuously varying composition will only form the basis for a predictive theory if link with environment. Our there is an explicit requires is a consequence sponse environmental gradient. realized differ. As reformulation that The the continuum re of two distinct types of and fundamental

for eco possibilities Their positive feature is that logical responses. and can be tested. Alterna they are operational tive theories also have this feature but lack several currently accepted tation which need features to be of plants and vege

(1972); statements, descriptive and speculative responses,

incorporated.

to these gradients of plants responses a consequence, patterns and vegetation mechanisms and types of com the physiological or interference) processes petitive (exploitative in the environmental will vary with position space defined by these gradients. of and the differences Our set of propositions 1, 2, 3) provide a set of multiple regarding these patterns. These need hypotheses to be tested in a variety of habitats and a much opinion (Tables fuller in the of where is needed knowledge are obtained. the results environmental space to a single habitat in Individual studies restricted a poorly specified part of the environmental space continue authors but to produce results. contradictory disturbance consider (Table 1) direct environmental ignore as temperature. of Comparison

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