1 Competition and Succession in Pastures –

Some Concepts and Questions

Philip G. Tow1 and Alec Lazenby2

1Department of Agronomy and Farming Systems, University of Adelaide, Roseworthy
Campus, Roseworthy, Australia; 263 Kitchener Street, Hughes, Australia

Background drastically reduced, largely because of cultivation for

cropping. The areas remaining generally owe their
Extensive observations made during the 19th and continuing existence to climatic, edaphic or topo-
early 20th centuries on the behaviour of plants graphic limitations to cropping or to their being set
growing together in communities led to the develop- aside as conservation areas (Fig. 1.6a, b, c). Species
ment of a number of principles on competition and originally characteristic of natural grasslands were well
successional changes (Clements et al., 1929). adapted to their environment. However, overgrazing
Sustained interest in interplant relations over the and deterioration in soil chemical and physical attrib-
past half-century has resulted in a large volume of utes have resulted in the degradation of many such
research and many theories on the subject. grasslands, and inferior plants, often regarded as
Publications such as Society for Experimental weeds, have replaced some of the original compo-
Biology (1961), Harper (1977), Wilson (1978), nents. Some degraded natural pastures are now being
Grace and Tilman (1990), Begon et al. (1996) and restored to conserve biodiversity and increase the
Radosevich et al. (1997) demonstrate both the attractiveness of the landscape (see Chapman,
progress made in understanding competition and Chapter 13, this volume). Effective rehabilitation and
succession and the inadequacies of such understand- appropriate management of such grasslands are each
ing, which is required for the optimum management dependent on a proper understanding of the princi-
and preservation of the world’s plant communities. ples involved in plant competition and succession.
This book is focused on competition and succes- Where climate, soil, landscape and financial
sion among plants in pastures; the term ‘pasture’ is incentives are favourable, large areas of improved
defined as vegetation used for grazing by domestic or pastures have been sown in many countries.
wild animals (Fig. 1.1a, b) and cutting by humans Cultivars selected and bred for high levels of pro-
for fodder conservation (Fig. 1.2). Grasses are a uni- ductivity, persistence and feed value are sown in the
versal component of such vegetation, which is thus most intensive farming systems as single species
often termed ‘grassland’. Legumes and other herbs (Fig. 1.7) or as mixtures of grasses or of grasses and
are other common pasture components (Fig. 1.3a, b, legumes (Fig. 1.3a, b). The principles of competi-
c). Shrubs and trees may also coexist with grasses tion and succession are directly relevant to the chal-
provided they are spaced widely enough to prevent lenge of maintaining desirable pasture composition,
crowding out of smaller plants (Figs 1.1b and 1.4). free of weeds, while achieving high levels of both
Naturally occurring grasslands (Fig. 1.5a, b), once productivity and utilization. These objectives
occupying vast areas of the world, have now been become even more difficult to achieve when the

© CAB International 2001. Competition and Succession in Pastures

(eds P.G. Tow and A. Lazenby) 1
2 P.G. Tow and A. Lazenby

(a) origin of the sown species is outside the country of

use – a fact which may add complexity to manage-
ment, to compensate for imperfect climatic and
edaphic adaptation. Conversely, similar complexity
arises from climatic change (see Campbell and
Hunt, Chapter 12, this volume) or from the effects
of change in land use on soil conditions (see
Chapman, Chapter 13, this volume). Because of
the many environmental and technological changes
currently affecting grassland and other plant com-
munities, an understanding of interplant and
plant–environment relationships is important for
interpreting, predicting and managing change in
(b) species composition.
The term ‘composition’ may be used in various
ways. For example, the term ‘botanical or species
composition’ can refer to: the presence of particular
species; a list of species present; or the proportion
(%) of various species in terms of plant numbers,
tiller or stem numbers, dry matter (DM) yield, leaf
area or ground cover. Botanical composition is
influenced by competitive relationships and may be
an indicator of a stage in succession.
The functioning and effects of competition and
succession in pastures differ from those in other
natural and man-made plant communities, such as
Fig. 1.1. (a) Sheep grazing in winter on a pasture of forests and annual crops. The most important of
annual medic (Medicago truncatula) in the such differences are caused by the grazing animal.
cereal–pasture zone of South Australia, Australia. Grazing and trampling reduce the height and some-
(b) A mix of grasses, shrubs and trees grazed by times the density of the canopy. This reduces com-
native animals in the Masai Mara National Reserve
petition for light between shoots and may
in Kenya.
indirectly reduce the intensity of root competition
if part of the root system reacts to defoliation by
growing more slowly or dying. Defoliation by graz-
ing and cutting reduces the competitive advantage
gained by plants which emerge earlier, have a larger
embryo to begin growth and have favourable attrib-
utes such as higher initial relative growth rates,
tillering rates, leaf expansion, root spread and
stature (Milthorpe, 1961).
Grazing of grasses encourages renewed growth
of existing tillers and development of new ones. In
mixtures, the timing and intensity of grazing affect
competitive relationships and resulting proportions
of the community components (Milthorpe, 1961).
Severe defoliation and trampling may kill existing
Fig. 1.2. A mixture of oats (Avena sativa) and
plants and leave gaps for invasion by others or result
common vetch (Vicia sativa) grown for silage at the
Roseworthy Campus of the University of Adelaide,
in soil erosion, perhaps on a large scale (see Davies,
South Australia, Australia. The oats are a strong Chapter 4; Kemp and King, Chapter 5; Wolfe and
competitor against vetch, but the legume can Dear, Chapter 7; Skarpe, Chapter 9; Peltzer and
compensate by climbing upwards with long, Wilson, Chapter 10; Garden and Bolger, Chapter
twining stems and tendrils. 11; and Chapman, Chapter 13, this volume).
 Competition and Succession in Pastures 3

(a)

(b)
Fig. 1.4. Vigorous spring growth of native grasses
following winter burning, in open Eucalyptus
woodland in the Undara Volcanic National Park in
far north Queensland, Australia.

Competition – Definitions and

Concepts
Definitions of plant competition and succession are
based largely on observation and experience or
(c) measurements of effects on plants, rather than on
an understanding of mechanisms (Tilman, 1990).
The following definition of competition proposed
by Clements et al. (1929) is still accepted by many:
Competition is a purely physical process. With few
exceptions, such as the crowding up of tuberous
plants when grown too close, an actual struggle
between competing plants never occurs. Competition
arises from the reaction of one plant upon the physical
factors about it and the effect of these modified
factors upon its competitors. When the immediate
supply of a single, necessary factor falls below the
Fig. 1.3. (a) A well-balanced mixture of perennial combined demands of the plants, competition begins.
ryegrass (Lolium perenne) and white clover (Trifolium
repens) in its third year, near Lutterworth, Leicestershire,
In Milthorpe’s (1961) words, the term competition
England. (b) A 4-year-old unirrigated pasture, describes ‘those events leading to the retardation in
composed principally of the perennial grass cocksfoot growth of a plant which arise from association with
(Dactylis glomerata) and the annual legume other plants. It results from the modification by
subterranean clover (Trifolium subterraneum), near Mt adjacent individuals of the local environment of
Barker in the Adelaide Hills region of South Australia. each particular individual’. This general definition
Each year, following opening autumn or winter rain, the could include such effects as allelopathy (the
legume must re-establish from seed in competition with adverse effects on one plant of a toxic substance
the established grass. (c) A balanced, irrigated pasture derived from another).
mixture of Grasslands Puna chicory (Cichorium
The following ‘working definition’, proposed by
intybus), WL516 lucerne (Medicago sativa) and Maru
phalaris (Phalaris aquatica), in late winter, at Dalby Begon et al. (1996), is applicable to pastures:
Agricultural College on the Darling Downs of southern ‘Competition is an interaction between individuals,
Queensland, Australia. Weeds have been excluded by brought about by a shared requirement for a
the dense, diverse mixture of sown species. resource in limited supply and leading to a reduction
4 P.G. Tow and A. Lazenby

(a) (a)

(b) (b)

Fig. 1.5. (a) A dense, vigorous stand of native (c)

bluegrass (Dichanthium sericeum) in central
Queensland, Australia. The originally vast areas of
such native grasses have been greatly reduced by
overgrazing and/or cultivation for cropping. (b) A
remnant of original prairie preserved in Texas, USA.

in the survivorship, growth and/or reproduction of

at least some of the competing individuals con-
cerned.’ However, the statement that competition is
an interaction between individuals is not a definition
in itself; competition is defined by the cause (limited
resources) and the net effect (yield reduction) of
Fig. 1.6. (a) Arid rangeland in Wadi Rum in the
competition. This illustrates the difficulty of provid- south of the Kingdom of Jordan. Such areas are
ing a precise definition of competition without a grazed periodically by the migratory sheep, goats
clear understanding of the mechanisms involved (see and camels of the Bedouin people. (b) Rangeland
Sackville Hamilton, Chapter 2, this volume). on uncultivated slopes near Digne, Alpes-de-Haute-
Goldberg (1990) has proposed a simple, mech- Provence, France. (c) Grassland in the Masai Mara
anistic framework for studying interactions National Reserve, Kenya, grazed periodically by
between plants, based on her observation that migratory herds of wild animals and domesticated
‘most interactions between individual plants actu- native cattle.
ally occur through some intermediary’. In the case
of competition for resources (e.g. plant nutrients changes in abundance of the resources. Plants can
and water) one or more of the competitors will be good competitors either by rapidly pre-empting
have an effect on the abundance of the resources and depleting a resource (by uptake) or by being
(intermediaries) and they will also respond to able to continue growth at depleted resource levels.
 Competition and Succession in Pastures 5

The depletion of light resources has sometimes

been measured in mixed-plant canopies (Stern and
Donald, 1962; Rhodes and Stern, 1978), but
depletion of soil water and nutrients by compo-
nents of mixtures is much more difficult to quan-
tify. Yet Tilman (1982, 1988, 1994) has based his
definition of competitive ability on the theory that,
over a number of years, the winning competitor is
the species (among those initially present) which is
able to reduce the concentration of the limiting
soil resources (e.g. available N) to the lowest level
and still maintain its population, i.e. it is the one
with the lowest resource requirement or R*. This
Fig. 1.7. A pure stand of the annual legume barrel mechanism of competition has been called the
medic (Medicago truncatula) at the Roseworthy
resource reduction model. The R* values for a
Campus of the University of Adelaide, South
group of species, if known, would predict the final
Australia. Such stands are kept as free of weeds as
possible, to maximize nitrogen input for livestock (equilibrium) outcome of competition among
feed and following crops, as well as for annual seed these species for a limiting resource; it should be
production. independent of the timing of establishment of
competing species, their starting proportions and
the initial sizes of individual plants. However, the
The latter case (classed as low net competitive experimental work supporting this definition has
response to competitors) may occur through con- been done in small, ungrazed plots. The resource
tinued uptake at low levels of resource, decreased reduction model may therefore be more applicable
resource loss from plant parts or increased effi- to lightly grazed, low-input grasslands, rather than
ciency of conversion of internal stores of the to intensively grazed pastures receiving regular,
resource to new growth (see also Peltzer and high inputs of nutrients (see also Kemp and King,
Wilson, Chapter 10, this volume). Chapter 5; and Peltzer and Wilson, Chapter 10,
The relative importance of above- and below- this volume).
ground competition has often been questioned. An alternative view of competition to that of
Milthorpe (1961) concluded from various experi- Tilman is that a plant will be competitively supe-
ments and observations on crops and pastures that rior if it has the capacity to capture (pre-empt)
‘competition between roots usually commences resources faster than others. This can be related to
long before the shoots are sufficiently developed to particular plant traits, such as high potential rela-
cause mutual shading’. Further, following an analy- tive growth rate (Grime, 1979). Plants that can tol-
sis of 23 competition studies, Wilson (1988) con- erate low levels of resource (e.g. plant nutrient)
cluded that root competition is usually more availability are classed as stress tolerators. It has
important than shoot competition in determining been hypothesized that differences among compet-
competitive balance between species. However, ing species in resource acquisition rates, once estab-
species vary in their response to root competition, lished, are maintained and magnified during
as found by Bolger (1998). He conducted an exper- competition because of positive feedback between
iment to compare the capacity of seedlings of a growth and resource capture (Harper, 1977; Grime,
number of southern Australian pasture plants to 1979; Keddy, 1990; Begon et al., 1996). This pro-
‘invade’ an established sward of phalaris, a peren- posed mechanism of resource competition has been
nial grass. Experimental treatments comprised vary- called resource pre-emption or asymmetric compe-
ing degrees of shoot and root competition from tition. It occurs, for example, when large plants
phalaris and varying levels of plant nutrient supply. intercept a disproportionate share of light, while
The species differed greatly both in the ability of small plants have very little effect on the light
their recruiting (intersown) seedlings to compete reaching the larger plants. In comparing the models
with established phalaris root systems and in their of Tilman and Grime, Wedin and Tilman (1993)
relative response to root and shoot competition (see explain that, while both of the above mechanisms
also Nurjaya and Tow, Chapter 3, this volume). of competition allow for an initial pre-emption of
6 P.G. Tow and A. Lazenby

resources by one species, Grime and Tilman differ grass was about 22% compared with a silvergrass-
on which mechanism determines the long-term free environment.
outcome of competition. Dominance of one species by another may also
Goldberg (1990) suggests that the two models occur in an established pasture as a result of a dif-
agree over a successional sequence which progresses ferential response to seasonal climate variation or to
from fast-growing species with rapid resource selective grazing or simply as a result of differences
uptake rates to slower-growing species that are tol- in growth habit. One of the tasks of grazing man-
erant of low resource levels. Tilman’s R* value for agement is to prevent excessive or prolonged domi-
species with the lowest resource requirement would nance of one desirable component of a pasture over
refer to dominant, highly stress-tolerant species in another, thus preventing excessive lowering of the
equilibrium (non-successional) communities. ‘presence’ of the latter (see Kemp and King,
Asymmetric competition may occur between Chapter 5; and Harris, Chapter 8, this volume).
plants of the same species (as part of intraspecific
competition) or of different species (interspecific
competition). It accounts for self-thinning, particu- Competition – Quantification of
larly in newly established pasture. Populations
experiencing the greatest degree of crowding (inten-
Effects
sity of competition) have the greatest size inequal-
In any practical consideration of competition in
ity, i.e. competition exaggerates underlying size
plant communities, it is of value to be able to quan-
inequalities (Begon et al., 1996). Thus self-thinning
tify the effect of competitive interactions on the
occurs in response to plant density, but the level of
components of the community and on the course
thinning is also modified by the availability of
of competition over time. In pasture communities,
resources, such as moisture and light.
it is useful if the results can be related to environ-
Plants which establish earliest not only have a
mental factors or to management treatments that
large adverse effect on later-appearing plants, but
have been applied. This should also lead to the
are themselves little affected by the latter. Thus the
defining of appropriate management for regulating
earliest-established plants tend to persist, while
interplant relations. Keddy (1989, 1990) has
attempts to invade their environs continue to fail,
defined competition intensity as ‘the combined
at least where the initial density of the earliest
(negative) effects of all neighbours on the perfor-
plants is high (see Fig. 1.3). This principle is used
mance of an individual or population’. It is mea-
where possible in pasture management to exclude
sured by comparing the performance of
weeds. However, it also means that the introduc-
components in a mixture with those in monocul-
tion of desirable species into existing swards is
ture, or comparing the performance of ‘target’
unlikely to succeed unless adequately sized gaps are
plants surrounded by neighbours with that of the
created by the use of cultivation, herbicides or
plants in plots cleared of neighbours. Grace (1995)
heavy grazing. Once gaps are created, the way is
has argued the inadequacy of using absolute differ-
open for rapidly establishing ‘opportunists’ (ruder-
ences between yields in monoculture and mixture as
als) to fill them. Gaps are also created and weeds
a measure of competition intensity. This is because
allowed to enter when desirable species die due to
the magnitude of the difference would depend not
extreme climate conditions or attack by insect pests
only on the relative competitive abilities, but also
and diseases. For example, the grassy weeds Vulpia
on the relative magnitude of monoculture yields.
spp. invaded large areas of southern Australian
Thus he proposed that a more appropriate index
lucerne-based pastures when Hunter River lucerne
would be one that reflected the proportional impact
(Medicago sativa cv. Hunter River) was decimated
of competition on plant performance, i.e.
by the spotted alfalfa aphid in 1978/79. Vulpia
then spread to other pasture areas as opportunity relative competitive intensity (RCI) =
arose. A survey of farms in south-eastern South performance in monoculture − performance in mixture
Australia and western Victoria showed that, by performance in monoculture
1998, Vulpia fasciculata (silvergrass) and other
Vulpia species were at a serious level of infestation De Wit and van den Bergh (1965) also pointed
on 1.8 million ha (Silvergrass Task Force, 1998). out that the intensity and course of competition
The average loss of gross farm income due to silver- between species in pasture could not be unambigu-
 Competition and Succession in Pastures 7

ously quantified by simply comparing the perfor- the course of competition, in place of grain yields
mance of the species in the mixture. First, yields of used for crops. The index to define the course of
individual species at particular times cannot be competition was called the relative replacement rate
equated with others, i.e. 1 g of one is not necessarily (ρ). Relative yields are used to define ρ of species a
the same as 1 g of another. They stressed the need for with respect to species b at the nth harvest with
a dimensionless measure, such as the relative yield respect to the mth harvest by
(yield of a species in mixture/yield of the species in n
ra / m r a
monoculture). They also pointed out that reference nm
ρ ab = n
to monoculture yields enables changes in growing rb / m r b
conditions and varying lengths of growing period to If ρ > 1, species a is the strongest competitor. If
be taken into account. If only differences between ρ < 1, species b is the strongest. If ρ is plotted on a
species in mixture are measured, it is difficult to logarithmic scale against time, the angle of the line
determine whether these are due to differences in with the horizontal is a measure of the relative rate
competitive ability or to differences in response to at which one species replaces another. The same
growing conditions. Including monoculture yields in course line may be obtained by plotting the ratio of
the formula helps to account for the latter. relative yields at successive harvests (van den Bergh,
An important measure of competition, devel- 1968). This course line is very useful for judging
oped by de Wit (1960, 1961) is the relative crowd- the direction of competitive relationships over time
ing coefficient (k) (see also Sackville Hamilton, but not for further quantitative analysis of the
Chapter 2, this volume). This is a measure of ‘com- mutual interference. Van den Bergh conducted
petitive power’, namely, the degree to which a experiments to show the effects on the course of
stronger competitor crowds a weaker one. De Wit competition of various factors, e.g. plant density,
studied numerous field experiments in which barley plant nutrient treatments and pH levels. De Wit
and oats were grown both in monoculture and in and van den Bergh also found that, almost invari-
mixtures, where various proportions of barley were ably, grass species were mutually exclusive
replaced by the same proportions of oats (replace- (RYT = 1), i.e. they were competing for the same
ment design). The results showed that, in the mix- resources and the relative replacement rate was
tures, one species always crowded the other out of independent of the relative frequency (sowing pro-
some of the space ‘allotted’ to it according to the portions) of the component species. This is an
composition of the sown mixture. Gains and losses important ecological concept. While not yet
were equivalent. Consequently, in terms of grain explained, it helps illustrate and predict how
yield, the relative crowding coefficient of barley (kb) stronger and weaker competitors interact when
with respect to oats was the reciprocal of the rela- competing for the same set of resources.
tive crowding coefficient of oats (ko) with respect to In contrast to the situation with mixtures of
barley, i.e. kb × ko = 1. Furthermore, the relative grasses, de Wit et al. (1966) found that a grass and a
yield total (RYT) = 1, where legume were not mutually exclusive when the
legume obtained N from symbiotic fixation. Their
grain yield barley in mixture
RYT = experiment was conducted with and without rhizo-
grain yield barley in monoculture bial inoculation of the legume. Without rhizobium
grain yield oats in mixture and N fixation, the grass and legume were mutually
+ exclusive (RYT = 1). With rhizobium, however, N
grain yield oats in monoculture
fixation gave the legume a competitive advantage.
In terms of competition theory, this means that the RYT was greater than 1 and the species were not
two species were crowding for the same ‘space’ or mutually exclusive because the legume had an addi-
resources. In these circumstances, yields of mixtures tional source of N not available to the grass. When
cannot exceed the yield of the highest-yielding course lines of the ratio of relative yields were
monoculture. drawn, over seven harvests, the lines of mixtures of
De Wit and van den Bergh (1965) and van den different sowing frequencies tended to converge and
Bergh (1968) showed that the above concepts also to approach equilibrium (no change, no one species
apply to mixtures of pasture grasses. They found winning competitively). These trends were attrib-
that yields of successive harvests provided an appro- uted to a combination of N fixation (which
priate measure of plant performance for defining favoured the legume competitively) and N transfer
8 P.G. Tow and A. Lazenby

from legume to grass (which favoured the grass dominance. Furthermore, white clover content in
competitively). It is now widely assumed that mix- pastures is also subject to long-term fluctuations or
tures of grasses and legumes, at least those based on cycles (Chapman et al., 1996).
white clover, have a capacity to regulate the N cycle Renewed interest in white clover–grass pastures
in the pasture (Chapman et al., 1996). over the past 20 years (reflected in the increased
In later experiments in both field and glasshouse number of relevant publications, for example, in
(Tow, 1993; Tow et al., 1997), trends with time in Grass and Forage Science) is related to the belief that
the ratio of relative yields provided further evidence clover N, compared with fertilizer N will reduce
of the tendency for a dynamic equilibrium to occur, costs, use of fossil energy and leaching of nitrate to
provided that: (i) one species did not remain domi- groundwater. Further, Ennik (1981, 1982), exam-
nant for too long; and (ii) growing conditions were ining experimental data in the literature, found
generally favourable to the growth of the legume. that the DM yield of a mixed white clover–grass
As indicated above, such course lines show if and sward receiving N fertilizer at varying levels was
under what conditions grasses and legumes tend always higher than that of a pure grass sward at the
towards equilibrium, but do not provide a means of same rate of mineral N application. This was
further analysing competitive interactions. A ten- because, with increasing application of N, the gain
dency for equilibrium should have a positive influ- in grass DM was higher than the loss of clover
ence on stability of botanical composition and DM. He also estimated that the amount of fertil-
species persistence. izer N needing to be applied to a pure grass sward
Where climatic conditions fluctuate over time, to obtain an N yield equal to that of a mixed sward
the course of competition may also fluctuate. This was about 80 kg N t−1 of clover in the mixture
may result in breakdown in equilibrium. However, (after the first tonne). This linear relationship,
the work of Tow and his colleagues quoted above accompanied by an inverse relationship between
provides evidence that, as long as dominance is not rate of fertilizer N input and clover content of the
too severe, there is a persistent tendency to equilib- mixture led to the conclusion that most of the fer-
rium. Equilibrium between species, or at least coex- tilizer N was taken up by the grass. Furthermore,
istence, is often said to be due to the fact that they he concluded that, while introduction of more
occupy different niches. In grass–legume mixtures, competitive clover varieties into a mixed pasture
the legume occupies a different niche in the sense may increase N yield of the mixture, it was unlikely
that it has an independent source of N. to increase DM yield.
The attainment of equilibrium or coexistence Improvement of clover yield, N2-fixation and
sometimes requires an input of management that persistence have all been recent objectives of plant
assists towards reducing the dominance of strong breeders, agronomists and modellers. (Caradus et
competitors, e.g. grass over legume. Such manage- al., 1996; Chapman et al., 1996; Evans et al., 1996;
ment is of most benefit if it achieves competitive Schwinning and Parsons, 1996). All agree that
balance without loss of productivity and with bene- effective production and utilization of grass–clover
fit to the grazing animal (see Davies, Chapter 4; pastures require understanding of the interactions
Kemp and King, Chapter 5; and Harris, Chapter 8, of the two components. This becomes all the more
this volume). important as attempts are made to achieve a combi-
Achieving a competitive balance is more com- nation of aims, such as: (i) increasing the yield of
plex than might be supposed. For instance, a gen- clover by breeding more competitive cultivars and
eral problem with white clover–grass pastures is the cultivars with a higher capacity for N2-fixation,
difficulty of maintaining the clover content of some while avoiding leakage of nitrate to groundwater;
30% thought to be desirable (Martin, 1960). This (ii) increasing total yield by the use of N fertilizer
might be simply a problem of reducing grass domi- without losing clover content; (iii) managing
nance by appropriate management. However, grass–clover swards for optimal animal production;
defining appropriate management of grass–legume and (iv) assessing new cultivars of white clover
competition and N relations has to take account of under grazing conditions (see also Nurjaya and Tow,
the spatial heterogeneity (patchiness) of clover con- Chapter 3; and Davies, Chapter 4, this volume).
tent brought about by spatially random urine depo- Experience with grass–clover mixtures provides a
sition. This keeps different areas in the field ‘out of reminder that competition usually operates in con-
phase’ with respect to surrounding grass or legume junction with other factors that affect companion
 Competition and Succession in Pastures 9

plants differentially. In such mixtures, the most with their competitive abilility. Such traits do not
important factors would probably be N2-fixation, N always define the mechanism involved, but they assist
transfer and selective grazing. Competition is some- in explaining or predicting competitive outcomes
times distinguished from ‘apparent competition’, (see Nurjaya and Tow, Chapter 3; Skarpe, Chapter 9;
where reduced yield of one component of a mixture and Peltzer and Wilson, Chapter 10, this volume).
may be due to differential effects of another organ-
ism on that component, e.g. selective grazing of
palatable species, leaving an unpalatable one in Succession
higher proportions; or the same effect by selective
attack by an insect pest. Begon et al. (1996) quote an Succession, the change in botanical composition
example discussed by Connell (1990) of an indirect over time, is currently a subject of great importance
effect of Artemesia bushes on the growth of associ- in both natural and sown pastures (as illustrated by
ated herbs. The beneficial effect of removing the the contents of this volume). Such importance
bushes on the growth of the herbs was initially attrib- arises because of the many changes that have
uted to reduced competition for water. It was then occurred over the past century, largely resulting
found that removal of Artemesia also discouraged from increasing intensification of pasture use.
deer, rodent and insect consumers of the herbs which Succession has long been linked to competition.
used this plant as a source of both food and shelter. More than 70 years ago, Clements et al. (1929)
Figure 1.8 and accompanying commentary illustrate concluded, from their North American research
just how complex interspecies relationships can be. and experience, that competition ‘is the controlling
In attempting to understand the mechanisms of function in successional development, and it is sec-
competition and to predict the outcome, many ondary only to the control of climate in the case of
researchers have identified morphological and physi- climaxes’. They also concluded that the regular out-
ological traits or characteristics of plants associated come of competition is dominance, the successful
competitors coming to control the habitat more or
less completely. Other components of the plant
community face suppression or even extinction.
As a feature of cyclic changes, Clements and his
colleagues envisage regular invasion of plant com-
munities from species outside. Hence their asser-
tion that:

The [successful] invading community is in harmony

with the changing climate, the one invaded is
correspondingly handicapped by it, and is all the more
readily replaced as a result of competition between
them. The course of events in edaphic habitats where
succession is occurring is much the same, but the
Fig. 1.8. Plants of the annual legume rose clover advantage to the invaders arises from the changes
(Trifolium hirtum) growing amongst a clump of a brought about by the occupants, which serve as a
perennial, native speargrass, near Bukkulla, progressive hindrance to possession.
northern New South Wales, Australia. The relation
between grass and legume may be quite complex. They then see the climax as the mature stage ‘in
In summer, the grass clump may intercept seed pods harmony with the climate’ and yet exhibiting an
of rose clover washed over the soil surface by heavy ‘annual departure in growth and numbers’, due to
storm rains. In winter, the grass is almost dormant climatic variation. In all these processes, Clements
and poorly competitive, when the clover establishes
et al. regarded competition as having a leading role
from seed and makes much of its growth. In spring,
as temperatures rise but soil moisture declines,
in determining the botanical structure of the vege-
plants of rose clover may survive only in the shelter tation. The competitive balance of various grass,
of speargrass clumps, as in the photo. However, it herb and shrub types in grassland is disturbed by
must set seed quickly before the new growth of variations in rainfall and is also ‘profoundly modi-
speargrass becomes too competitive. fied by grazing, burning or cutting’.
10 P.G. Tow and A. Lazenby

These and related conclusions were subse- invasion, effects of climate change and an increas-
quently translated into a successional approach to ing range of technological inputs. The so-called
rangeland management (the so-called range succes- state and transition model seems to satisfy these
sion model) and a practical system of range classifi- needs. It involves the concept of ‘thresholds of envi-
cation (Westoby et al., 1989; Laycock, 1991). As ronmental change’, which cause ‘transition’ from
summarized by Westoby et al. (1989): one discrete or stable ‘state’ of the vegetation to
another. Such transition requires the imposition of
the [range succession] model supposes that a given a threshold of stress or perturbation. The prediction
rangeland has a single, persistent state (the climax) in or early detection of an impending threshold would
the absence of grazing. Succession towards this climax allow management action to be taken to maintain
is a steady process. Grazing pressure produces changes
or achieve desirable botanical structure and produc-
which are also progressive and are in the opposite
direction to the successional tendency. Therefore the
tivity levels.
grazing pressure can be made equal and opposite to Westoby et al. (1989) suggest that, for the effec-
the successional tendency, producing an equilibrium tive use of the state and transition model, recorded
in the vegetation at a set stocking rate. information on particular areas of rangeland should
include catalogues of possible alternative states,
The main tool of range management for the range possible transition pathways, opportunities for pos-
succession model is thus the level of stocking rate. itive management action and hazards which may
However, ‘vegetation changes in response to grazing produce an unfavourable transition. The experi-
have been found to be not continuous, not mental testing of hypotheses (e.g. opportunistically
reversible or not consistent’, particularly in arid and during the occurrence of isolated events or
semi-arid areas. These observations have led to a sequences of events) should be a regular feature of
general questioning of the range succession model. information gathering. This needs to be accompa-
In recent years, the need for an alternative nied by the estimation of probabilities of occur-
model to describe and assess rangeland condition rence of climatic circumstances relevant to
and dynamics has been discussed by many workers, particular transitions. Such information should also
e.g. Westoby et al. (1989), Friedel (1991), Laycock be of value for describing and managing other types
(1991) and Humphreys (1997). Particularly ques- of grassland, at least for long-term pastures.
tioned has been the need to manage rangeland to Similarly, the proposals of Friedel (1991) should be
achieve a single, climax state or at least some desir- applicable to a wide range of grasslands and pas-
able, stable state in equilibrium with an economic tures. She argues for the need to monitor botanical
stocking rate. A ‘stable’ system (in terms of botani- composition and yield of arid and semi-arid range-
cal composition) returns to the original steady state lands in order to detect the approach of a ‘thresh-
after being disturbed or deflected. Some researchers old’ of change from one state to another. She
and practitioners prefer a system to have ‘resilience’, presents evidence that this is feasible from monitor-
namely, the capacity to adapt to change, without ing programmes and the use of multivariate analy-
necessarily reverting to the original state. What is ses and ordination techniques. The research
regarded as important would depend on both the suggests that rangeland which is deteriorating may
economic and conservation goals of management retain the capacity to recover up to a certain point,
and the opportunities and limitations set by the beyond which it cannot readily return to its former
environment and available technology. Rangeland state. Some factor, such as drought, fire or flooding,
stability and resilience may each be important in usually coincides with excessive grazing to ‘tip the
particular situations and can be envisaged as depen- balance’. Appropriate monitoring needs to be com-
dent to some extent on interspecific competition. bined with an understanding of plant–environment
The above authors thus favour a model of relations to allow prediction of approaching thresh-
rangeland dynamics that caters for the occurrence olds, thereby enabling preventive action to be
of multiple states of vegetation structure, changing taken.
influences on these states and the need for flexibil- The role of competition in determining vegeta-
ity of short-term aims and management. A model tion structure and succession has received little crit-
of this nature should also be appropriate for many ical attention in the above debate. It may be that, in
other grasslands, where botanical structure has been arid and semi-arid areas, the overriding influences
or is being greatly modified by over-grazing, weed on plant community structure and succession are
 Competition and Succession in Pastures 11

management (e.g. stocking rate effects) and peri-

odic climatic events. If so, competition may play a
lesser role than that claimed by Clements et al.
(1929). It may have an increasing effect on pasture
plant community structure with increasing rainfall
and the accompanying greater plant density and/or
productivity. The role of competition may also vary
with the level of soil nutrient availability. These
suppositions have long occupied the attention of
ecologists and continue to do so (see Peltzer and
Wilson, Chapter 10, this volume). A clear under-
standing of such matters is needed for the effective
use of a state and transition approach to pasture
management, with its need for clear definition of Fig. 1.9. The annual legume Medicago truncatula
establishing from self-sown seed in early winter,
thresholds of change and transitions between rela-
amongst the stubble of the previous season’s cereal
tively stable states. Indeed, its effectiveness as a
crop. This illustrates a phase in the traditional South
model will depend on an understanding of all rele- Australian ley farming system.
vant plant–environment, plant–plant and
plant–animal relationships. These must be under-
stood at the scale of both individual plants and the encourages and frees the manager to use a wide
wider ecosystem. range of information and management options to
As mentioned earlier in this chapter, the regula- achieve goals of pasture production, botanical com-
tion of dominance and invasion by highly competi- position and long-term stability or resilience.
tive weeds is important in managing succession or Management based simply on opportunism is no
retaining a desirable stable or resilient botanical basis for achieving long-term pasture outcomes.
structure (see Kemp and King, Chapter 5; and Wolfe What is needed, for all types of pastures, is a system
and Dear, Chapter 7, this volume). The manage- developed within a framework of clearly defined,
ment problems posed by the heterogeneity of vegeta- long-term goals and an understanding of factors
tion and associated environments in rangelands and affecting competition and succession. Management
pastures, together with current concerns about loss decisions, incorporating various pathways and time
of species diversity, are encouraging increasing inves- periods, could then be taken to achieve the goals.
tigation on these topics (see Clark, Chapter 6; and
Chapman, Chapter 13, this volume).
In farming situations, a different form of com- Conclusions
plexity is provided by rotations of pastures with
crops. For example, in the so-called South Ecologists and agronomists are currently making
Australian ley farming system (Fig. 1.9), weeds of considerable efforts both to overcome and prevent
the annual legume-based pastures, if allowed to set degradation and to improve the long-term perfor-
seed, are likely to re-appear in and compete mance of natural and sown pastures. Achieving
strongly with following crops (see Trenbath, these objectives requires an understanding of the
Chapter 14, this volume). One principle that can processes involved, e.g. processes leading to decline
be used in changing competitive relationships and in productivity, loss of valuable species and ingress
botanical composition involves sowing mixtures of of weeds, as well as those associated with improved
pasture species, which together, at appropriate den- productivity and stability. Plant competition is an
sities, are more competitive against weeds than the important factor controlling these processes.
legume alone (see Figs 1.2 and 1.3c; see also Harris, However, the nature of such competition is not yet
Chapter 8, this volume). fully understood. There are other factors – abiotic
One attitude engendered by adoption of the and biotic – which can have a major effect on the
state and transition model is that range manage- conditions under which competition occurs.
ment can be seen as a continuing ‘game’, the object Parallel with research on the mechanisms of
of which is to seize opportunities and evade hazards competition is work that has produced indices to
as far as possible (Westoby et al., 1989). This measure outcomes of plant interactions. These
12 P.G. Tow and A. Lazenby

indices enable comparisons to be made of the mining the botanical composition of pastures
effects of genotype and environmental factors on remains unclear. Even so, integrating the bank of
such outcomes. Other investigations have resulted information and improved understanding arising
in the description and classification of the botanical from the array of ecological and agronomic research
structure of grasslands and any changes over time, should provide a real opportunity to develop man-
thereby providing a basis for management deci- agement systems to achieve long-term use and sta-
sions. Yet the precise role of competition in deter- bility of grasslands.

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