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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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35
A new model
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
phenomena'
36 Whittaker These were propositions in eucalypt forest in The evidence for the continuum
(1972).
positive new hypotheses or new, more robust In this paper, we attempt frameworks. to re-express the continuum theory of vegetation robust detected
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
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
example,
vegetation
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
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
communities
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,
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
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
individualist optima
to the optimal
along an environmental
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.
and
continuum
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
New
model
for a Continuum
Environmental Three
types
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
resource size
such
as food
inclusive The
response
of a limiting that due to toxicity at levels beyond under natural conditions normally experienced decline
(Fig. 3a).
3. Direct gradients for plants are those that have
plants
herbivory
1978). Ellenberg suggested that plant species have similar optima and physiological response curves on the basis of his early experiments (Ellenberg
? 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
niche
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.).
(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
the ability growth (i. e., survive) under resource availability (Parsons
literature
fundamental
40
unlimited tional
and we combinations
need
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
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
curve is modified to response curve observed realized response of competition, and conditions herbivory
disease. Experiments
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
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
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
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
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
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
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.
% DOMINANCE
cost
and
absolute
amount
of e.g. low
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
(1980)
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.
patterns
of species
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
Ellenberg
(1953,
1954; Mueller-Dombois
regarding (1979) concerning species
&
pri
response
43 in vegetation, of for survival mary strategies nutrient-ratios Tilman 1982) (Tilman regarding
workers
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
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
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
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
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
of variety apparent, contrary to his theoretical assumptions. on patterns of There is no common agreement richness species along gradients (Table 3). Grubb
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
alternative
Species response
environment
Ellenberg
optima shape
As
for resource
gradient
Not considered
Not considered
Not
considered
Not
considered
Not
considered
Not
considered
Tilman
Law Nested
of diminishing
returns,
Not
considered
Bazzaz
Variable Nested
shape,
Various
Limited
range
Similar
Optima,
optima, shape,
Limited range
Certain combinations non-viable of
responses
45
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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
forms
particular C4 plants)
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
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
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
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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