CH 7

C&S in Pastures Chap 07 22/10/01 1:38 pm Page 119
7 The Population Dynamics of Pastures,

with Particular Reference to Southern Australia
E.C. Wolfe1 and B.S. Dear2

1School of Agriculture, Charles Sturt University, Wagga Wagga, New South Wales,
Australia; 2NSW Agriculture, Wagga Wagga, New South Wales, Australia
Introduction the dynamics of plant communities. Then, in a

series of case-studies, pasture species/varieties that
In Australia, as in other continents, agricultural have been notably successful in Australian agricul-
development has resulted in substantial changes to ture will be related to these models, and evaluated
grassland ecosystems. Australian agriculture has in terms of the processes and principles that have
passed through several distinct phases since white underpinned their competitive success. In a con-
settlement in 1788 (Table 7.1; Shaw, 1990; Barr cluding section, past experience and experimenta-
and Cary, 1992). In the 19th century, an initial tion will be considered in relation to the future
exploration phase was supplanted by exploitation of integration of conservation and agriculture in
the Australian landscape for grazing and crop produc- Australian rural lands.
tion, and then followed periods of consolidation,
amelioration and restoration during the 20th century.
Not surprisingly, in response to this development, The Impact of Agricultural
profound changes occurred in the botanical composi- Development on the Pastures of
tion of Australian grasslands (Moore, 1970). Among Southern Australia
the original native species and among those species
that were either accidentally or deliberately intro- Early in the 19th century, the occupation of rural
duced, there have been notable failures and survivors. Australia by British immigrants proceeded slowly
In this chapter, a brief historical assessment will but, by 1860, driven in part by the discovery of gold
be given of the impact of development on grass- in 1851, land settlement encompassed the south-
lands in the agricultural areas of southern Australia, eastern quarter of Australia from north of Adelaide
which are characterized by Mediterranean-type to beyond Brisbane, as well as part of western
(south-western, southern) and temperate (south- Australia (Shaw, 1990). In 1862, New South Wales
eastern) climates; these grasslands have a minimum (NSW) and Victoria were both supporting about
mean annual rainfall of about 300 mm. Examples 6 million sheep, with Queensland only a little
of the botanical changes that occurred in response behind. There were many difficulties during this
to grazing, plant introduction and fertilization on exploration phase (Shaw, 1990; Barr and Cary,
the tablelands (predominantly non-arable, grazing) 1992), but the availability of suitable grasslands was
and slopes/plains (arable, farming) will be explored not one of them (Barr and Cary, 1992).
in relation to three conceptual models that have According to Shaw (1990), the period between
been used, worldwide, to describe and explain the 1850 and 1890 marked the heyday of the pastoral-
interplay of climatic, edaphic and biotic factors on ists, who developed huge grazing properties with
© CAB International 2001. Competition and Succession in Pastures

(eds P.G. Tow and A. Lazenby) 119
120 E.C. Wolfe and B.S. Dear
Table 7.1. Agricultural development in Australia, 1820–2000.a
Livestock numbers ( 106)

Wheat area
Phase Year Sheep Cattle (ha 106)
Exploration 1820 0.3 < 0.1 0.01

phase 1842 6 < 0.1 0.06
1851 17 0.2 0.1
Exploitation 1861 21 3.8 0.3
phase 1871 40 4.3 0.5
1881 65 8.0 1.2
1891 106 11.1 1.3
Consolidation 1901 72 8.5 2.1
phase 1911 97 11.8 3.0
1921 86 14.4 3.9
1931 111 12.3 6.0
1941 125 13.6 4.9
1950/51 116 15.2 4.2
Amelioration 1960/61 153 17.3 6.0
phase 1970/71 172 24.4 6.5
Restoration 1980/81 131 25.2 11.3
phase 1990/91 162 23.6 9.2
1996/97 120 26.8 10.9
aThe statistics for 1820–1950/51 were taken from Shaw (1990) and the more
recent values from ABARE (Knopke et al., 1995) and the Australian Bureau of
Statistics.
axes, dams, artesian bores, fences and better tech- railway network, in NSW. A common problem was
niques of sheep breeding and husbandry. By 1891 the decline of cereal yields on land that had been
there were 62 million sheep in NSW, 20 million in farmed for several years (Donald, 1967).
Queensland and 13 million in Victoria (see Table However, not all of the 19th century was char-
7.1); between 1860 and 1894, the Australian sheep acterized by exploitation. The rapid expansion of
population had risen from about 20 million to 100 mechanization and transportation from 1870 and
million and cattle from 4 million to more than 12 the establishment of agricultural colleges and
million (Shaw, 1990). experimental farms during the 1890s were notable
A combination of factors, notably overstocking, developments in terms of their immediate benefit
the invasion of pastoral lands by rabbits introduced and future impact.
in 1859, a general economic depression and bank As outlined by Barr and Cary (1992), the
collapses in the 1890s, and a severe drought from southern Australian wheat industry was rescued
1895 to 1902, arrested and then reversed the from decline at the turn of the century by the new
pastoral boom. Erosion, pasture degradation and a techniques of dry farming, purposeful wheat breed-
fall in livestock numbers were the consequences, ing and superphosphate fertilizer. However, crop
outcomes predicted by P.E. de Strzelecki. In an yields on the poor Australian soils were still low by
1840 report to Governor Gipps on his travels to the world standards and bare fallowing both depleted
Australian Alps and Gippsland, Strzelecki expressed the soil of organic matter and rendered it liable to
his concern about the exploitative practices of over- wind erosion. Green manuring with oats or lucerne
grazing and burning (Hancock, 1972). Meanwhile, was advocated, but it was not until the 1930s that a
the farming of wheat, barley and oats had expanded technique of ley farming1 with annual pasture
steadily in area (see Table 7.1), first in South legumes was developed and promoted (Puckridge
Australia (Meinig, 1954) and Victoria (Barr and and French, 1983). Subterranean clover (Trifolium
Cary, 1992) and then, with the development of a subterraneum L.), which had been discovered and
Population Dynamics of Pastures 121
promoted unsuccessfully by Amos Howard in the fertilized annually more than doubled between
1890s, was the basis of the ley farming system 1950 (7 million ha) and 1973, when a superphos-
introduced at Rutherglen Research Station in phate subsidy was removed. These improved pastures
north-eastern Victoria (Barr and Cary, 1992). Ley consisted mainly of subterranean clover, annual
farming with annual medics (Medicago spp.) was in medics, lucerne (Medicago sativa L.) and, in the
use in the 1930s at the Roseworthy Agricultural higher-rainfall areas (> 600 mm) of NSW, Victoria
College in South Australia (Callaghan, 1935). and Tasmania, perennial grasses, such as phalaris,
Fertilized ley pastures, by setting in train a new perennial ryegrass (Lolium perenne L.) and cocks-
succession of botanical changes in response to foot (Dactylis glomerata L.).
added fertilizer and N2-fixation, had the potential Finally, during the 1970s, 1980s and 1990s, a
to alter markedly the nature of grazed pastures in reappraisal of pasture development took place. This
the farming zones. This topic will be dealt with reappraisal occurred in response to: (i) a worsening
later in this chapter. Because of the depression and cost-price squeeze (Gruen, 1990); (ii) an apparent
then war, there was little change in on-farm prac- widespread decline in the productivity of pasture
tices or outputs between 1930 and 1950 (Donald, legumes following the occurrence of new disease,
1967; Gruen, 1990), and land degradation contin- pest and weed problems (Gramshaw et al., 1989);
ued. However, investments in agricultural research and (iii) evidence of widespread land degradation
during this consolidation phase produced some phenomena, such as eucalyptus dieback, soil acidifi-
notable discoveries that were to underpin the rapid cation and salinization (Goldney and Bauer, 1998).
expansion of improved pastures from 1950 to 1970 The area of sown pastures in Australia has remained
on both arable and non-arable sites. These discover- static at around 27 million ha since 1970, with
ies included successful searches (mainly within declines from 1970 to 1985 in the area fertilized
Australia (Cocks et al., 1980)) for new varieties of and the rate of fertilizer applied (Gramshaw et al.,
subterranean clover that were intermediate between 1989). Since then, problems of oversupply in the
the mid-season variety Mount Barker (found in wool industry and changing community attitudes
South Australia, commercialized in 1906) and the towards conservation have given rise to initiatives
early strain Dwalganup (commercialized in western that aim to integrate conservation and agricultural
Australia, 1929); a strain (cv. Hannaford) of annual production, with the objective of achieving sustain-
medic (barrel medic, Medicago truncatula Gaertn.) able production systems. This restoration phase
that was commercialized in South Australia in (Goldney and Bauer, 1998) has been marked by: a
1938; the development as a sown species of a strain reduction in the sheep population (see Table 7.1);
of Phalaris tuberosa L. (syn. aquatica), a perennial
better documentation of the effects of and solutions
grass that was capable of surviving summer drought
to the environmental problems created by agriculture;
(Oram and Culvenor, 1994); the realization that
programmes and legislation to protect areas of
deficiencies of phosphorus and sulphur were wide-
native grasslands and woodlands; a shift in emphasis
spread in Australian soils, many of which also
from action at the farm level to the catchment level;
needed one or more of the minor (trace) elements
and improved partnerships between scientists,
copper, zinc, molybdenum, manganese, iron and
farmers and the community.
boron (Williams and Andrew, 1970); elucidation of
the legume – Rhizobium symbiosis (Williams and
Andrew, 1970); and the development of a virulent
strain of the myxomatosis virus for rabbit control Pasture Dynamics During the
(described by Barr and Cary, 1992). Phases of Agricultural
The above knowledge, allied with the 1950s Development
boom in wool prices, financial incentives for invest-
ment in agriculture (Gruen, 1990) and the advent Models of pasture dynamics
of aerial agriculture (Campbell, 1992), ushered in
an amelioration phase in temperate Australian There are at least three different conceptual models
grasslands. According to the account of this phase that have been used as a framework for describing
by Crofts (1997), the area of sown pastures and explaining the interplay of climatic, edaphic
increased from around 5 million ha in 1950 to in and biotic factors on the dynamics of pasture com-
excess of 25 million ha by 1970: the pasture area munities and for managing these communities. The
first two of these models, the Clementsian theory of Bolger, Chapter 11, this volume). This pathway of
succession and the state and transition model, were change in plant communities, from a pre-settlement,
recently discussed fully by Humphreys (1997); climax (stable) vegetative state through an unstable
these models are illustrated and explained in a continuum of several disclimax stages, where stability
following section. A third theory, the was more or less maintained by ‘management’, was
competition–stress–disturbance (CSD) model, was consistent with the original linear succession model
developed and used by Grime (1977) to classify first proposed by F.E. Clements in 1916
plants according to the combination of characteris- (Humphreys, 1997). In terms of Clements’s theory,
tics they display in response to three primary eco- the activities of fire, grazing, clearing and fencing
logical factors. These factors are: (i) competition opposed the natural successional tendency towards
(with the high vegetative competitive ability of pristine, climax grassland.
some species accounting for their dominance); (ii) Loss of grazing-susceptible plant species, open-
stress (with certain plant species adapted to and tol- ing the sward to native and exotic invading species,
erating unproductive conditions); and (iii) distur- nutrient redistribution and changes in the seasonal
bance (with some species adapted to grazing extraction and replenishment of soil water are the
disturbance, and with ruderal species possessing an processes that presumably influenced the outcome
ability to invade and grow in severely disturbed but of plant competition and the succession of plant
potentially productive environments). All three the- communities in Australian grasslands, over time
ories – succession, state and transition, and CSD – and space. While the main catalyst for the botanical
have some usefulness in interpreting the changes in changes that took place was grazing (a disturbance
vegetation that have occurred in natural and factor, in terms of Grime’s CSD model), the
improved pastures in southern Australia. evidence for its specific effects is largely anecdotal.
Early reports made during the exploration and
exploitation phases (Barr and Cary, 1992) indicated
Species changes during the exploration and that kangaroo grass was abundant and palatable to
exploitation phases livestock. Kangaroo grass was sensitive either to
defoliation or treading or both; it did not persist
Moore (1970) outlined the changes in pasture wherever sheep or cattle were grazed (Moore,
species that occurred over several decades in typical 1970). Subsequently, it was shown in South Africa
Eucalyptus woodland – grassland communities on (O’Connor, 1996) that the persistence of T. trian-
the slopes and tablelands of southern NSW in dra depended on lax defoliation, which enhanced
response to clearing, higher grazing pressures (from both the recruitment and the survival of seedlings.
sheep, cattle and rabbits) and the application of Several other native perennial grasses were
superphosphate fertilizer. The original climax vege- protected in part from grazing by mechanisms such
tation, dominated by tall, warm-season, perennial as less palatable herbage (for example, Bothriochloa
tussock grasses, such as kangaroo grass (Themeda ambigua S.T. Blake, red grass) or spiny seeds (Stipa
triandra Forsskal), plains grass (Stipa aristiglumis F. spp., spear or corkscrew grasses). On the northern
Muell.) and poa tussock (Poa labillarderi Steud.), slopes of NSW, three-awned speargrass (Aristida
was presumably well adjusted to the ebb and flow ramosa R.Br.), an unpalatable species, became
of the native herbivores (kangaroos, wallabies, bird co-dominant with red grass on extensive areas of
life) and occasional fires. Once sheep and cattle lightly grazed grasslands (Williams, 1979).
were introduced to the tablelands and slopes in the However, it has subsequently been shown that A.
1830s–1840s, accompanied by timber-clearing ramosa is sensitive to defoliation and the balance
operations, there began a sequential progression can be shifted back towards more palatable species,
(Fig. 7.1) in the botanical composition of the grass- notably wallaby grass (Danthonia spp.) and subter-
lands, towards a disclimax community (or, more ranean clover, by heavy grazing with flocks of sheep
correctly, a number of disclimax communities). applied strategically in summer–autumn, coinciding
Such communities contained an array of grazing- with the flowering and seedling establishment of
tolerant, cool- and warm-season native grasses, the speargrass (Lodge and Whalley, 1985).
together with various naturalized annual grasses Johnston (1996) noted that the causal relation-
and forbs that had been introduced into Australia ship between grazing (defoliation, trampling) and
in agricultural seeds and feeds (see also Garden and grass species composition has not been seriously
C&S in Pastures Chap 07 2/11/01 8:47 am Page 123
Original climax community

Tall warm-season perennial grasses
Themeda triandra
Stipa aristiglumis
Poa caespitosa
grazing
Disclimax community
Short cool-season perennial grasses
Danthonia spp.
Stipa falcata
more intense grazing
Dwarf cool-season perennials + dwarf warm-season perennials

Danthonia spp.
Stipa falcata, Chloris truncata
further grazing
Dwarf cool-season perennials + cool-season Mediterranean annuals

Danthonia
(now polymorpha)*
Vulpia spp.*, *
*, *, a*
Panicum effusum, Stipa falcata, Chloris truncata
phosphorus addition and grazing
Current pasture ley community with sown species,

introduced cool-season annuals + warm-season annuals
Trr *, *
*, Vulpia spp.*, Lolium rigidum *
Echium plantagineum *, *
*
Arctotheca calendull *
Fig. 7.1. Botanical changes in pasture in the cropping zone of south-eastern Australia, based on Moore (1970)
and modified by Dear (1998). Asterisks denote naturalized species, to distinguish them from native species.
challenged. He listed several factors that may have as well as timber clearing and reduced fire frequency.
been involved in the replacement of the original According to Johnston (1996), the persistence to
native grasses with other grasses; these include the present day of certain productive and nutritious
changes in plant–soil water and nutrient relations, C3 native grasses, notably weeping meadow grass
(Microlaena stipoides (Labill.) R.Br.) and the wal- to the grazing factor, superphosphate application
laby grasses, may be due, at least in part, to their and N2-fixation by legumes became important
preference for shade and/or the complementarity of agents in determining the direction and pathways
their growth cycle and phenology to the original C4 of botanical change. Both benefits and problems
grasses, which are taller and better adapted to dry (Table 7.2) were initiated by pasture improvement,
habitats and hot seasons. and this period of rapid change was followed by
Other important components of the degraded one of reassessment. Consequently, the pathways of
grasslands included many grasses, forbs and botanical change in pasture communities became
legumes that became naturalized after their entry to numerous and complex.
Australia. Sometimes this naturalization followed Humphreys (1997) postulates many reasons
their accidental introduction in agricultural produce why the dynamics of grasslands are better described
from various destinations along the sea routes from by a ‘state and transition’ model instead of
Europe and the Mediterranean region to Australia Clements’s succession model. The state and transi-
(for example, Vulpia spp., silver grass, Hordeum lep- tion model is able to represent a set of multiple
orinum (Link), barley grass, and Arctotheca calendula pathways and vegetation states occurring in
(L.), Levyns, capeweed). In other cases, plants may response to several sets of factors. In addition, it is
have been introduced deliberately as ornamentals flexible in accommodating the notion of resilience
(for example, Oxalis pes-caprae L., soursob, Silybum (ability to recover) in an ecological system, whereas
marianum (L.) Gaertn., variegated thistle, and Clements’s theory is too focused on the concept of
Echium plantagineum (L.), Paterson’s curse or a single climax state and on grazing management as
‘Salvation Jane’ (Michael, 1970)). Most of the the dominant factor that drives succession.
strains of subterranean clover found in suburbs of In Fig. 7.2, adapted from Lodge and Whalley
Perth by J.S. Gladstones (1966) entered Australia (1989) and Garden et al. (1996), the state and tran-
(accidentally, not deliberately) before 1870, and sition model is used in an attempt to summarize
some annual Medicago spp. were naturalized by the the nature and timing of the main changes that
early 1900s (Crawford et al., 1989). have occurred in grasslands on the tablelands of
Michael (1970) attributed the competitive success NSW, particularly during the last half-century. At
of these alien species to their adaptability to disturbed one extreme of the time–management continuum,
environments and/or the absence of their native pests native and naturalized pastures were ploughed up,
and competitors. Why they were so successful in sown with perennial grasses and clovers and fertil-
competition with the native perennial grasses is open ized; initially, such pastures were dominated by the
to speculation. A possible reason is that the winter- sown legumes until soil N levels from rhizobial
growing Mediterranean annuals depleted soil water in activity were sufficient to allow the grasses (and
spring such that the summer-growing perennials were weeds) to compete effectively with the legumes.
unable to survive the long, dry summers. Another The botanical progression from initial legume
factor may have been selective grazing of the perenni- dominance towards eventual grass (or other non-
als by sheep over summer, thereby reducing the legume) dominance was dependent on a minimum
above-ground green material of the perennials and annual rate of superphosphate application (to stim-
presumably lowering energy reserves in their roots ulate and favour legume growth) and grazing (to
and crowns. In the case of annual legumes, one clear enhance the transfer of fixed N to the associated
advantage was their capacity to fix N and thrive in species). For example, in grass–white clover pas-
the low-nitrogen (N) soils of southern Australia. tures on the northern tablelands of NSW, the
clover-dominant phase was intensified, but the
onset of grass dominance was hastened following
Instability of grazed pasture communities high annual rates of superphosphate (375 kg ha1)
over the last half-century instead of intermediate (125–188 kg ha1 year1)
rates (Wolfe and Lazenby, 1973). The cessation of
During the last 50 years (the amelioration and superphosphate application to perennial
restoration phases), the dynamics of pastures ryegrass–white clover pastures in the same locality
became even more complex in response to develop- resulted in a loss of the sown species and a corre-
ments that accompanied or followed the pasture sponding increase in the proportion of the native
improvement revolution (1950–1970). In addition red grass (Cook et al., 1978).
Table 7.2. An updated list of the factors that are, or may be, associated with the decline of pasture
legumes in southern Australia.
Medic decline, South Australia (Carter et al., 1982)

Reduced spraying to control insect pests of pastures (earth mites and lucerne flea)
Reduced application of superphosphate fertilizer to pasture
Spread of sitona weevil
Increased cropping intensity and consequent grazing pressure
Poor grazing management and fodder conservation practices
Increased use of herbicides in the cropping phase of the rotation
Reduced undersowing of medic into cereal crops
Rapid spread of pasture aphids (spotted alfalfa aphid, blue-green aphid, pea aphid) after introduction
in 1977/78
Increased use of nitrogen fertilizers on crops (less need for medic ley)
Apathy and despondency concerning value of medics
Medic decline, South Australia (Denton and Bellotti, 1996)
Drought
Sulphonylurea herbicides (suspected)
Higher populations of root lesion nematodes
Zinc deficiency
Rhizoctonia solani
Subterranean clover decline, southern slopes, NSW (Hochmann et al., 1990)
Suboptimal supply of phosphorus
Root rot associated with Phytophthora clandestina
Soil acidity
White clover decline, northern tablelands, NSW (Hutchinson et al., 1995)
Climatic stress, particularly the effect of January–March (late summer) rainfall on stolon survival
Set-stocking at high rates, encouraging the presence of competitive annuals
Soil nitrogen (N) build-up (of secondary importance)
Annual legumes, Australia (Gramshaw et al., 1989)
Economic trends (lower returns for beef and wool compared with cropping, removal of fertilizer
subsidies, farm cost inflation during the 1970s)
Insect pests and diseases (lucerne pathogens, clover scorch, root rots of Trifolium spp., effects of
spotted alfalfa and blue-green aphids on lucerne, medics and other pasture legumes)
Land degradation (soil acidity, salinization, compaction)
Changes in crop production techniques (more frequent cropping, longer cropping sequences, partial
substitution of N fertilizers and pulses for pasture legumes)
High-rainfall zone and annual zone pastures, Australia (Wilson and Simpson, 1993)
No regular and standardized surveys of the state of pastures, but ad hoc surveys have indicated
suboptimal clover content and the significant presence of unsown annual grasses, particularly in
legume leys in the cropping zone
Quantitative evidence is lacking on both the extent and the nature of change in botanical composition
and productivity
Factors that are claimed to be associated with low legume content, such as stocking rate, climatic
variation, pasture age and management, are listed and reviewed. The list is similar to that of Carter et
al. (1982) at the top of this table
At the other extreme of the management con- unless the guidelines developed by Campbell
tinuum, large areas of native pastures on the NSW (1992) were used. These guidelines were to sow the
tablelands were aerially sown with subterranean grasses when rainfall is likely to be effective, use
clover and treated with superphosphate from time herbicide to minimize competition from the exist-
to time. Introduced perennial grasses were usually ing vegetation and minimize seed theft by ants. In
not sown from the air; if they were, their establish- contrast to the difficulties associated with grass
ment was frequently unsuccessful (Wolfe, 1968) establishment, the establishment of aerially sown
C&S in Pastures Chap 07
% Legume
126
Annual biomass production
(kg green DM ha–1 year–1
ORIGINAL VARIOUS STATES
STATE
Native 2500 0
+G
Original Modified
1850 native Native/naturalized
native/
22/10/01
pastures naturalized
pastures Naturalized
3000 –
5000 10
Degraded
+G –F
1:38 pm
+F Well managed
E.C. Wolfe and B.S. Dear

+G
–F Improved +F Degraded by acidity,

weedy or salinity
Page 126
Modified 3500 –
1900 native thistly 8000 20
pastures +F +F Improved, weedy
+F
+G +D
Improved Improved
Improved +F, E ryegrass or +F grass
clover cocksfoot dominant,
dominant dominant less stable
Modified +F, L (–E) +E
1950 native/ 5000 –
naturalized Improved 9000 30
pastures clover Improved
dominant +E phalaris +F Improved
(+E) dominant phalaris
dominant,
+F stable
+F, L, E
1960 1980 2000
Fig. 7.2. The application of the state and transition model to the generalized pathways of botanical change that have occurred in permanent pastures, south eastern
Australia. The boxes are ‘states’, and arrows represent the general direction of transition due to the main agents of change (G, grazing; F, fertilizer; L, legumes; E,
exotic grasses; D, drought). Depending on the circumstances, a transition may or may not be reversible.
legumes (subterranean clover and/or white clover) and white clover); the apparently accidental intro-
was generally satisfactory. Further, so long as phos- duction of three pasture aphids (the spotted alfalfa
phate applications were maintained, legumes aphid (Therioaphis trifolii (Morrell) f. maculator),
became dominant (Willoughby, 1954) until the the blue-green aphid (Acyrthosiphon kondoi Shinji)
pastures were invaded by N-loving species, such as and the pea aphid (Acyrthosiphon pisum Harris))
annual grasses and forbs (Rossiter, 1964) or thistles into Australia in the late 1970s, seriously reducing
(Michael, 1970; see also Fig. 7.2). Many such the productivity and persistence of lucerne and
species were more aggressive for moisture, light and annual medic pastures (Panetta et al., 1993); and
nutrients and/or were better able to exploit the soil the development of new races of plant pathogens,
conditions. Thus, many of the native perennial such as Phytophthora clandestina races that attack
grasses were suppressed or eliminated by competi- subterranean clover (Dear et al., 1993b).
tion or by high grazing pressure. In summary, both progressive and episodic
There were several intermediate scenarios between occurrences produced in Australian grasslands a
these extremes. For example, less palatable tussocky range of pasture states (see Fig. 7.2). The potential
grass species, such as poa tussock (P. labillarderi and actual changes, including those that occurred
Steud.), were present in low numbers on the southern before the amelioration phase, are represented
tablelands of NSW. They increased in frequency and adequately by the state and transition model, which
size over time in response to the N fixed by the intro- conforms fully with the ecological principle of suc-
duced legumes (Fisher, 1972); then, heavy grazing cession (progressive change) but which avoids the
pressure on the palatable species in the gaps between inadequacies (linearity, inflexibility) that are
the tussocks exacerbated tussock dominance, response embodied in the concept of a linear succession
to fertilizer declined and the swards deteriorated into towards a monoclimax (Humphreys, 1997).
unproductive, N-deficient grasslands. Much the same
sequence occurred on parts of the north-western
slopes of NSW, where three-awned spear-grass Ley pastures in the cropping belt
became dominant (Williams, 1979), and on the
central tablelands, where the aggressive and indi- Ley pastures in Australian croplands (Puckridge
gestible serrated tussock (Nassella trichotoma (Nees) and French, 1983) represent a different situation
Arech.) colonized tracts of non-arable country from that of the permanent pastures used for grazing
(Campbell, 1998). This latter species, introduced sheep and cattle on the coast and tablelands of
from South America, became a serious weed of the south-eastern Australia. In such leys, the opportu-
grasslands of South Africa and New Zealand, as well nity is available to resow pasture legumes, and the
as Australia (McLaren et al., 1998). time frame of the pasture phase between crop cycles
On many soil types, the prolific growth of pasture is short (typically 1–5 years).
legumes that was evident in the early years of fertil- A useful conceptual representation of the popula-
ized pastures in most areas of southern Australia did tion dynamics of the species in ley pastures is
not continue. There were several reasons for this. Grime’s (1977) triangular CSD model (Fig. 7.3) of
From the 1970s, the general malaise of ‘pasture the competition, stress and disturbance factors that
legume decline’ (Carter et al., 1982) was evident, influence the distribution and abundance of plant
together with progressively developing incidence of species. As noted by McIvor (1993), pasture and
specific problems, such as soil acidity (Cregan and grassland species are located near the centre of the
Scott, 1998), and the occurrence of certain pests CSD triangle; competition (C), stress (S) and
and diseases of pastures (Panetta et al., 1993). Some disturbance (D) are all important but none is over-
of the possible biological, nutritional, economic whelming. The annual legumes used in leys are
and social factors involved in pasture decline are ruderal (R) species that tolerate disturbance.
listed in Table 7.2. Ruderal species (C-R, S-R or C-S-R plants with short
Many of the environmental phenomena leading life cycles, high reproductive effort or other strategies
to change were episodic rather than progressive. for dealing with disturbed sites) take advantage of dis-
Examples of episodic events include severe turbance to colonize; they are favoured in productive
droughts, which occurred in most decades (leading croplands, which are frequently disturbed by crop
to the loss of drought-susceptible, introduced phases (cultivation, herbicide application) and by
perennials, such as perennial ryegrass, cocksfoot grazing during the pasture phase.
100 0
75 C 25
Di
n
(% etitio
stu (%)
rba
)
mp
50 50
nc
Co
e
C–S C–R
C–S–R
25 75
S S–R R
0 100
100 75 50 25 0
(%)
Stress
Fig. 7.3. Adapted from Grime’s (1977) competition–stress–disturbance model representing the factors of
competition, stress and disturbance on plants, and the location of plants that possess primary strategies (C,
competitors; S, stress-tolerators; R, ruderal species that are adapted to disturbance) and secondary strategies
(e.g. C–S, C–S–R) of adaptation to these influences.
At the other extremes of Grime’s model, stress- of profuse seeding and several have dormancy or
tolerators (S – low growth rates and low reproductive other seed conservation mechanisms and/or they are
effort) are favoured on less disturbed, less productive adapted in some way to grazing). A list of some com-
locations (e.g. acid soils). In contrast, competitors (C mon components of ley pastures and an overview of
– high growth rate, low reproductive effort) are their CSD tolerance are given in Table 7.3. The
favoured in undisturbed, productive situations (such population dynamics of subterranean clover and its
as abandoned cropland or lightly grazed pastures). In companion species will be discussed in a later section.
permanent pastures or long leys in Australian Problems have been reported in the ley pasture
agriculture, pasture species that exemplify success are system in Australia. They are due in part to
tolerant of stress (for example, drought) and distur- landowners reacting to higher economic returns
bance (grazing) and/or, in favourable locations, are from crops than from livestock (Reeves and Ewing,
efficient competitors for light and nutrients. 1993) and to the apparent decline in the legume
Examples of such plants are phalaris, lucerne and, in content of ley pastures (Carter et al., 1982;
locations with high annual rainfall (> 750 mm), Hochmann et al., 1990; Gramshaw et al., 1989).
perennial ryegrass. Some of the reports of legume decline and the factors
However, in any community, the CSD balance that may be or are associated with it are listed in
varies both spatially and temporally (McIvor, 1993). Table 7.2. Solutions have been found in liming
This is particularly so in pasture leys where the (Cregan and Scott, 1998), improved agronomy
annual species are examples of C–S–R plants, which (Hochmann et al., 1990) and the selection and
are adapted variously to competition (for moisture, release of tolerant cultivars of annual legumes
light and nutrients), stress (acid soils, seasonal water- (Collins and Stern, 1987). A promising technique –
logging) and disturbance (ruderal species are capable winter cleaning with herbicides (a combination of
Table 7.3. Characteristics of some common temperate pasture species in relation to their tolerance of
competition (C), stress (S) and disturbance (D).
Species Characteristics
Subterranean clover C May be outcompeted by annual grasses for P and for light; subclover
Trifolium subterraneum effective in suppressing some weeds in their rosette stage (for example,
St John’s wort)
S Tolerant of Al and Mn toxicities in acid soils; yanninicum subspecies tolerates
waterlogging; brachycalycinum subspecies is adapted to neutral–alkaline soils
D Outstanding tolerance of grazing – seedlings unattractive to livestock, plus
defoliation in winter stimulates inflorescence production, burr burial and
seed yield; prolific seed production, and effective seed conservation mech-
anisms (embryo dormancy, hard seed)
Annual medics S More effective than subclover in tolerating water stress and Zn deficiency,
Medicago spp. but sensitive to acid soils
D High levels of hard seed guarantee persistence in semi-arid localities
Serradella S Very tolerant of Al toxicity in acid soils, deep rooting allows it to access Zn
Ornithopus compressus at depth in sandy soils
D Prefers sandy soils, coverage of seed by drifting sand is important in the
breakdown of hard seeds
Balansa clover S Tolerant of waterlogging
T. michelianum D High levels of hard seed
Persian clover S Tolerant of waterlogging, heavy-textured soils
T. resupinatum D Prolific seeder, high levels of hard seed
Lucerne C Deep-rooted, crown habit – competes well for water
M. sativa S Some cultivars susceptible to insects, root and crown rots
D Adapted to rotational grazing
Perennial ryegrass C Competes well for light with white clover but shallow-rooted habit means
Lolium perenne susceptibility to Australian droughts
D Free-tillering, recovers well from grazing; will re-establish well from seed
Phalaris C Allelopathic, most persistent of the introduced perennial grasses due to
Phalaris aquatica deep rooting habit
S Susceptible to soil acidity
D Strong tuber, resists overgrazing
Annual ryegrass C Competes well for light and nutrients with clover, crops
Lolium rigidum S Very tolerant of Al and Mn toxicities in acid soils
D Prolific seeding, with dormancy mechanisms
Barley grass C Effective scavenger for soil P
Hordeum leporinum D Prolific seeder, seeds unattractive to livestock
Silvergrass C Allelopathic
Vulpia spp. D Herbage unattractive to livestock; prolific seeder
Capeweed D Mature plants unattractive to livestock; prolific seeding and complex seed
Arctotheca calendula dormancy mechanisms
Paterson’s curse C Tall when mature, shading out competitors
Echium plantagineum D Not eaten by cattle and horses at any stage, or by sheep when mature
Saffron thistle D Prolific seeding; herbage not attractive to livestock
Carthamus lanatus
Skeleton weed S Deep-rooting and therefore drought-tolerant; susceptible to biocontrol agents
Chondrilla juncea D Prolific seeding
simazine and paraquat) – has been developed to component of leys (Peoples et al., 1998) and the
reduce weed content and boost the legume content adoption of pasture monitoring protocols to help
of ley pastures (Thorn, 1992). Further improve- farm managers identify key indicators of pasture
ments in the effectiveness of legumes in ley pastures legume production and to guide management
are likely through the increased use of lucerne as a accordingly (Crosby et al., 1993; Paul, 1999).
However, there are additional threats, such as the derived from an Australian temperate pastures data-
evolution or introduction of new weed ecotypes or base (Pearson et al., 1997). The current distribution
new disease races and the development of herbicide of introduced pasture species is broadly depicted in
resistance in weeds, that are likely to cause problems Fig. 7.4 (after Moore, 1970; Hill, 1996). The main
in legume leys. The failure of Group A and Group B features of this map are inland aridity barriers to
selective herbicides to control annual ryegrass the occurrence of all species, together with other
(Lolium rigidum Gaudin), a common component of boundaries (climatic, soil pH) that are applicable to
wheat-belt pastures before the advent of these selective the distribution of annual legumes (Donald, 1970).
herbicides, is one issue that is now causing particular However, despite the emphasis in Australian agron-
concern (Powles et al., 1997). Another concern with omy on exotic pasture species, native pastures still
herbicides is the deleterious effect of some residual cover a much larger area than that of introduced
herbicides on pasture legumes, particularly on alka- plants (see Table 7.5). While pasture legumes must
line soils, where the degradation of these herbicides is be based on the locally adapted variants of exotic
slow (Gillett and Holloway, 1996). germ-plasm, there is a strong case for the use of
management options that utilize adapted native
grasses and for the inclusion of a wider range of
Summary perennial grasses, particularly C4 species, for pas-
toral use (Johnston et al., 1999). Johnston et al.
During the last two centuries, there have been (1999) questioned the amount of research and
profound shifts in the botanical composition of development effort that has gone into replacing
temperate pastures in Australia, and the pace of indigenous grasses with exotic introductions, many
change has increased in recent decades. Because of of which fail to persist over the frequent droughts
the complexity of the interactions between pasture that characterize the Australian climate. They
species and environmental phenomena, no one reviewed the evidence for the persistence, produc-
model is available to explain the dynamics of tivity and nutritive value of several species of native
pastures. However, the state and transition model is grasses in grazed pastures, and argued for strategies
a useful and flexible way of representing and under- that utilize adapted, palatable grasses, such as wal-
standing pasture dynamics. The focus of the CSD laby grass, in low-input situations, or even C4
model is on the short-term response of individual native grasses in areas that are prone to hydrological
plant species to these three factors and thus it com- imbalance. Importantly, Johnston et al. (1999)
plements the plant community focus of the state acknowledged the folly of a ‘one-or-the-other’ phi-
and transition model. The use of these two models losophy, compared with an approach that achieves
is recommended as a way of understanding both complementarity between a low-input, conservative
the ecology and agronomy of grasslands/pastures. approach to pasture management and the high-
They are used in the next section in case-studies input, exotic approach to pasture improvement,
that illustrate the success of some pasture species in which has produced notable gains in the produc-
the Australian environment. tivity of Australian agriculture.
In the winter-dominant rainfall areas of south-
ern NSW and northern Victoria, the presence of a
Successful Australian Pasture persistent perennial grass in permanent, grazed
Species pastures appears essential for several reasons.
Annual pastures in such areas have a potential mid-
Pasture components – exotic or native? winter imbalance in the supply (relatively high) and
demand (low) for water and nitrate in the soil pro-
A recent estimate (Hill, 1996) of the potential areas file (Johnston et al., 1999). A persistent perennial
of adaptation in Australia for the main sown pasture component increases the pre-winter soil water
species – all of which were originally introduced deficit (for example, 135 mm for annuals, 210 mm
into Australia – is given in Table 7.4. This vast for perennials (Whitfield, 1998)) and thereby helps
potential may be compared with another recent overcome this potential imbalance. Minimizing the
estimate, that of the relative importance of pasture flows of water beyond the plant roots counters dry-
types in NSW (total area of sown pastures 5–6 land salinity (Passioura and Ridley, 1998) and
million ha; Australia 25–30 million ha) (Table 7.5), reduces the rate of soil acidification (Ridley et al.,
Table 7.4. Estimated potential areas of adaptation for nine temperate annual and perennial
pasture species for Australia (Hill, 1996).
Pasture species Area (million ha) % of freehold + leasehold land
South-eastern Australia
Subterranean clover 60.8 67.8
Balansa clover 23.7 26.5
Persian clover 30.7 34.3
Barrel medic 27.0 30.1
Serradella 37.4 41.7
White clover 20.7 23.0
Lucerne 86.4 96.4
Perennial ryegrass 19.4 21.6
Phalaris 33.9 37.8
Total, south-eastern 89.6 100.0
South-western Australia
Subterranean clover 20.8 94.0
Balansa clover 4.9 22.3
Persian clover 5.8 26.0
Serradella 7.2 35.6
Barrel medic 17.1 77.3
White clover 0.2 1.0
Lucerne 9.3 42.1
Perennial ryegrass < 0.2 < 1.0
Phalaris 2.7 12.0
Total, south-western 22.1 100.0
Table 7.5. The relative importance (on an area basis) of pas-

ture species, genera and types in NSW, 1995 (from Pearson et
al., 1997).
Description Percentage by area
Unimproved native pasture 62.5

Improved (fertilized) native pasture 16.8
Subterranean clover 14.8
White clover 6.0
Lucerne 4.5
Phalaris 3.4
Cocksfoot 2.2
Annual medics 2.1
Perennial ryegrass 1.5
Tall fescue 1.1
Annual ryegrasses 0.4
Serradella 0.2
1999). Such differences are meaningful to the in the soil profile, poorly buffered soils) (Passioura
achievement of hydrological balance, particularly and Ridley, 1998).
between the 600 and 800 mm average annual rain- The better water use of palatable perennial
fall isohyets in northern Victoria and southern grasses enhances livestock production compared
NSW. Soil degradation is common in these areas with pastures based on annuals or poorly persistent
due to a combination of factors (recharge, salt loads perennials (Axelsen and Morley, 1968). Furthermore,
LIMITS TO ADAPTATION (TEMPERATE SPECIES)

A = ARID BOUNDARY L
W = WARM BOUNDARY EMERALD
C = COLD BOUNDARY
pH = pH BOUNDARY (Medics)
SC,BM P
WC,PR
BM
A
GERALDTON
A ARMIDALE
SC,BM A pH
L MINNIPA
SC,BM CONDOBOLIN
L L
P ESPERANCE
P BM pH WAGGA
KANGAROO HORSHAM
PERENNIALS & ANNUALS ISLAND
P C
SC
ANNUALS WC, PR
SC = SUBTERRANEAN CLOVER SC
BM = BARREL MEDIC
PERENNIALS INLAND
L = LUCERNE LIMITS
C
P = PHALARIS
WC = WHITE CLOVER & PR = PERENNIAL RYEGRASS
Fig. 7.4. The limits to the growth and survival of pasture species (after Moore, 1970; Hill, 1996).
a persistent perennial grass component in pasture Phalaris – a persistent grass in Australia

prevents or reduces the incidence of undesirable
The ‘Australian’ commercial cultivar of phalaris is,
weeds, such as thistles (Michael, 1970), and the
according to experiments (e.g. Hill, 1985) and
ingress of unpalatable native and introduced tussock
grasses, as discussed earlier. experience (Watson, 1993), the most persistent of
the temperate perennial grasses that have been
introduced and sown in Australia. In southern
Case-studies NSW and Victoria, varieties of phalaris are capable
of reliable persistence down to a median annual
Some case-studies of successfully introduced pasture rainfall of 500 mm, whereas perennial ryegrass will
plants are reviewed below. Successful pasture species in persist only where the rainfall is at least 700 mm.
Australia are those which, in our environment, are Between these isohyets, where phalaris and, to a
stable (resistant to change), resilient (able to recover) lesser extent, cocksfoot are the only reliable perennial
and productive. Case-studies of a number of intro- grasses, there is a considerable area of land (see Fig.
duced pasture plants that have been successful in the 7.4) at risk due to land degradation. Even in wetter
Australian environment are relevant to the future ideal tableland localities where more than 700–800 mm
of sustainable production. They demonstrate some rainfall is received annually, phalaris has survived
principles that apply to the success or failure of desir- occasional droughts that have killed cocksfoot, tall
able pasture species and highlight lessons that should fescue (Festuca arundinacea Schreb) and perennial
have been learnt. The dynamics of phalaris swards and ryegrass (FitzGerald et al., 1995).
subterranean clover pastures are treated in most detail, A deep-rooted and prostrate habit, dense tiller-
reflecting their success and greater importance in ing, partial summer dormancy and the presence of
Australian agriculture than is the case elsewhere. a large underground rhizome are features associated
with the outstanding persistence of the original the plant. There are also some qualifications that
‘Australian’ strain of phalaris (Oram and Culvenor, need to be made to the use of phalaris to create
1994). The persistence of ‘Australian’ phalaris and relatively stable improved pastures.
the importance of stocking rate was illustrated by First, the newer, winter-active cultivars of
Hutchinson (1992), who compared over 28 years phalaris released during the 1960s and subsequently
the presence of phalaris with annual grasses in pas- (cvs Sirocco, Sirolan, Sirosa) have been found to be
tures grazed by sheep on the northern tablelands of up to 50% less persistent, measured as basal cover,
NSW (Fig. 7.5). The stability and resilience of than the ‘Australian’ phalaris cultivar (Culvenor and
phalaris following grazing and drought, at least at Oram, 1992). According to Oram and Culvenor
low (ten sheep ha1) and medium (20 sheep ha1 (1994), ‘Australian’ phalaris has a more prostrate,
in earlier years, then 15 sheep ha1) stocking rates densely tillered habit than the newer, winter-active
(see Fig. 7.5), contrasts with the death of white cultivars (Sirosa, Sirolan), and it has more capacity
clover (Hutchinson et al., 1995; Hutchinson and for rhizomatous spread; for these reasons it is suited
King, 1999), perennial ryegrass, tall fescue and to heavy grazing. Selection in grazed swards for basal
cocksfoot in improved pastures on the northern cover, which has a higher heritability than the
tablelands in droughts such as those that occurred spreading ability of spaced plants (Oram and
in 1965, 1980–1982 and 1994 (FitzGerald et al., Culvenor, 1994), is a potential way of improving
1995). Similar observations on the susceptibility of the persistence of future cultivars of phalaris.
perennial grasses to high stocking rates and drought Secondly, as reported in Oram and Culvenor
have been made in long-term grazing experiments (1994), phalaris was found to be intolerant of high
and on farm paddocks on the central tablelands soil aluminium (Al) levels, which, together with a
(Kemp and Dowling, 1991) and the southern high availability of soil manganese, may adversely
tablelands (Axelsen and Morley, 1968) of NSW. affect plants in highly acid soils (pHCa < 5.2)
The ecological importance of a relatively stable (Cregan and Scott, 1998). While a build-up in soil
perennial grass component and the high cost of Al levels may be constrained by the ability of well-
re-establishing this component if it is lost make it established, deep-rooted phalaris to capture leached
important that persistence, rather than ‘productiv- nitrate, attempts to establish phalaris on soils that
ity’, be the first priority in selecting grass cultivars. are already acid may be thwarted by the sensitivity
Many farmers are reluctant to grow phalaris to Al of the root growth of the establishing phalaris
(Barr and Cary, 1992) because of perceivable but seedlings, particularly if phalaris is competing with a
avoidable management problems (palatability, companion weed species that is tolerant of high
dominance, animal disorders) and other beliefs acidity. The effect of differential tolerance to Al on
(establishment difficulty) associated with growing competitive relationships in perennial grass–annual
Low stocking Medium stocking High stocking

25 25 25
Phalaris – basal cover (%)
20 Stable node 20 20
Stable node
15 15 15
High
resilience No
10 10 10 resilience
Low 1980–1982
resilience
5 5 5
1965 1980–1982 1965 1980–1982
Droughts Droughts 1965
0 0 0
0 6 12 18 24 30 0 6 12 18 24 30 0 6 12 18 24 30
Other grasses – basal cover (%) Other grasses – basal cover (%) Other grasses – basal cover (%)
Fig. 7.5. The stability and resilience of phalaris from 1964 to 1991 (inclusive) on the northern tablelands of
NSW (after Hutchinson, 1992).
ryegrass binary mixtures 24–36 weeks after sowing balance between their strong competitive ability
was shown in a recent pot experiment (Rubzen, under conservative stocking versus their vulnerabil-
1996; Table 7.6). The grasses used were phalaris (cv. ity to increased levels of disturbance from the
Sirosa, sensitive to Al) and cocksfoot (cv. Porto, combination of high stocking rates and grazing
highly tolerant), and the ryegrass ecotypes were preference’. Stresses, such as drought and poor fer-
either collected from an acid soil site where Al was tilizer management (Cook et al., 1978; Hutchinson
present at potentially toxic levels (A-ryegrass) or and King, 1999), exacerbate this vulnerability.
from an alkaline soil site that contained no free alu-
minium (B-ryegrass). In the alkaline soil pots, both
Lucerne, and competition between the
ryegrasses suppressed phalaris or cocksfoot, but in
components of lucerne mixtures
acid soil the competitive outcomes were determined
by the level of tolerance of each of the grasses to In southern Australia, lucerne (M. sativa L.) has an
aluminium: the A-ryegrass ecotype suppressed the area of potential adaptation that is greater than that
sensitive phalaris but A-ryegrass was suppressed by for any other pasture species (Hill, 1996; see also
the highly Al-tolerant cocksfoot cultivar. These results Fig. 7.4). In practice, the wider adoption of lucerne
indicate that the difficulty of establishing a sensitive has been constrained by its susceptibility to acid
plant type on acid soils may be exacerbated by the soils, the need for a rotational grazing regime to
presence of an adapted competitor. Sources of ensure its survival (Leach, 1978; Lodge, 1991), the
aluminium tolerance for phalaris are currently being incidence of insect pests and disease (Lodge, 1991)
used in a breeding programme to produce a tolerant and the attitudes of farmers and graziers, who per-
cultivar (Oram and Culvenor, 1994). ceived lucerne to be a special-purpose pasture rather
In summary, phalaris is the most persistent of than an all-rounder. Hence, lucerne was usually sown
the perennial species introduced into Australian in ‘hay paddocks’ as the only component of a pasture.
agriculture. However, due to the constant distur- However, there is advocacy of lucerne as a compo-
bance of grazing, successional change in perennial nent of permanent or semi-permanent grass–clover
pastures is consistently directed towards annualiza- pastures and with subterranean clover or medic in ley
tion (Hutchinson and King, 1999). According to pastures, as a means of enhancing livestock produc-
Hutchinson and King (1999), the loss of sown and tion (Wolfe et al., 1980), N2-fixation (Peoples et al.,
palatable perennial grasses can be explained ‘as a 1998) and/or water extraction from the soil profile
Table 7.6. Relative changes in the yielda of the componentsb of 50 : 50 binary mixtures
(replacement series design) compared with the yield of each component in monoculture
(Rubzen, 1996).
Soil treatment in pot Binary mixtures
A-ryegrass/phalaris B-ryegrass/phalaris
Acid soil +87% > 82% 50% < +39%
(pH 3.7, 2.3 g ml Al) A-ryegrass/cocksfoot B-ryegrass/cocksfoot
37% < +40% 85% < +86%
A-ryegrass/phalaris B-ryegrass/phalaris
Alkaline soil +73% > 70% +69% > 65%
(pH 8.2, 0.01 g ml Al) A-ryegrass/cocksfoot B-ryegrass/cocksfoot
+68% > 77% +72% > 67%
a Regrowth measured from the second to the third harvest, i.e. from 24 to 36 weeks after
sowing.
50 value = 272 M, very highly tolerant
b Grasses were cocksfoot (cv. Porto): ED Al c
phalaris (cv. Sirosa): ED Al50 value = 28 M, sensitive

Ryegrasses were A-ryegrass: ED Al50 value = 188 M, highly tolerant
B-ryegrass: ED Al50 value = 21 M, highly susceptible
c ED Al
50 value is the equivalent dose (concentration) of aluminium in solution that would
produce a 50% toxic response (in this case, a 50% reduction in root extension compared
with the nil-effect control).
(Crawford and Macfarlane, 1995; Dear, 1998). This with livestock (Collins, 1978). During summer, the
potential warrants a short consideration of the pool of hard seed (seed with an impermeable testa)
dynamics of lucerne mixtures. is partially protected from grazing by burr (pod)
In permanent pastures, lucerne is not a particu- burial, but losses of up to 50% of the seed bank
larly compatible component when associated with a over summer have been recorded (Dear and
perennial grass. Either the grass competes strongly for Jenkins, 1992). During winter, when grazing pressure
moisture with lucerne during summer and autumn is high (supply of green herbage < demand for green
(Wolfe and Southwood, 1980), or the rotational feed), the continued close grazing of the prostrate
grazing management that is essential for the plant’s herbage stimulates seed production in spring
survival (McKinney, 1974; Leach, 1978) does not (Collins, 1978). During spring, low grazing pressure
suit the growth/survival of the companion grass. on green herbage (supply > demand) and burr burial
In ley pastures, too, there is incompatibility protect the developing inflorescences. The only time
between the components of the recommended when subterranean clover is vulnerable to grazing is at
lucerne–annual legume mixtures. Annual legumes, the seedling stage; fortunately, livestock avoid grazing
such as subterranean clover, are usually grown with the young seedlings.
lucerne to provide winter feed when lucerne growth In spite of the adaptation of subterranean clover
is poor. These lucerne mixtures are frequently to grazing, pastures commonly contain a number of
unstable (Leach, 1978) and, in lower-rainfall environ- other vigorous sown and invading species, with the
ments (<500 mm annual rainfall), gaps develop sown clover component comprising 30% or less of
between the lucerne plants. The gaps are either bare the pasture biomass. The nature of competition
or, when soil N levels increase, occupied by oppor- between subterranean clover and other species and
tunistic annual grasses, such as Vulpia spp. Surface soil between strains of subterranean clover is important
erosion is encouraged, along with high levels of dust from several perspectives. These include the selection
contamination in the wool. Wolfe and Southwood of cultivars that are more vigorous and successful in
(1980) found that lucerne grew best with Geraldton, environments where clover decline is occurring and
the least productive of three subterranean clovers used, the need to replace oestrogenic strains (high-
and that seed yield of subterranean clover could be formononetin strains, which cause ewe infertility)
increased by increasing the lucerne row spacing. More with low-formononetin cultivars. An account of the
recently, it has been shown that lucerne can reduce key population characteristics that determine the
subterranean clover persistence in two ways: reducing success of the species provides an ideal introduction
the seed set of the subclover and reducing the to a consideration of the relationships of subter-
establishment of clover seedlings in autumn. Lucerne ranean clover with its competitors.
dries out the soil surface rapidly during clover seedling Rossiter (1966) reviewed the characteristics that
germination and establishment, particularly when seemed to be associated with the long-term success
there is an early autumn break and temperatures and of strains of subterranean clover, namely their sur-
evaporation rates are high (Dear and Cocks, 1997). vival/productivity over a number of seasons or
This results in a smaller proportion of the seed pool crop/pasture cycles. He listed the seed-producing
successfully establishing (Dear, 1998). capacity of a strain as an important determinant of
Hence, while lucerne is a valuable pasture species, success or failure. However, it is our experience
its use has been somewhat constrained, due in part to with testing hundreds of subterranean clover strains
its need for rotational grazing and to its incompatibil- and crossbreds over several years in NSW that,
ity with other pasture components. The improvement rather than inherent differences between strains in
of relationships between lucerne and companion their potential seed production, success differs only
species is a worthwhile topic for future research. according to how well the fit is between their inflores-
cence development (triggered by the temperature and
light regime (Archer et al., 1987)) and the soil mois-
The success of subterranean clover and other
ture profile in spring. For maximum seed production
components of ley pastures
of subclover (similar to annual medic, about 200 g
m2 (Wolfe, 1985)), moisture conditions must be
CHARACTERISTICS OF SUBTERRANEAN CLOVER. In non-limiting during the flowering and seed-develop-
Mediterranean-type climates, subterranean clover is ment phases (Collins, 1981; Blumenthal and Ison,
a species that is well adapted to continuous grazing 1993), a total of about 70 days. This ideal is rarely
achieved in the Australian environment, and a tar- ber of seedlings that survive to well below the target
get seed production of 60 g m2 is sufficient for a of about 1500 plants m2 that is needed to opti-
high-quality subclover pasture (Dear et al., 1993a). mize the early dry-matter production of subclover.
A practical guideline for optimizing the balance Hence, in most environments, moderate to high
between spring herbage production (later-maturing levels of residual hard seededness (20–60%) are
cultivars produce more spring herbage) and the sought in subclover cultivars, even in mid-season
amount of seed is to choose a cultivar that flowers strains (Dear et al., 1993a).
and sets seed just before soil moisture usually runs Increased levels of hard-seededness can also be
out in spring. used to guard against a failure to set adequate levels
The strong relationship between rainfall and of seed due to herbicide damage, insect attack or
time to maturity of subterranean clover observed in drought (Dear and Sandral, 1997; Fig. 7.6).
environments of southern Australia does not always Indeed, near the arid boundary for annual legumes,
hold in the plant’s countries of origin. Piano et al. such as at Condobolin in central NSW (Cornish,
(1993) found that, although mean populations of 1985), annual medics are preferred to subterranean
T. subterraneum subsp. subterraneum collected in clover – not only because they are more tolerant of
Sicily increased with increasing rainfall, in T. subter- water stress during seed development (Wolfe,
raneum subsp. brachycalycinum the effect of altitude 1985), but also because cultivars with very high
was greater and that of rainfall less marked. There is levels of residual hard-seededness are available
also evidence that seed-production characteristics of (Cocks et al., 1980). On the other hand, in cool,
subterranean clover strains of similar maturity can moist environments, such as northern NSW
vary. Piano and Pecetti (1997) found that, despite (Lodge et al., 1990) and Tasmania (Evans and Hall,
Geraldton and Seaton Park commencing flowering 1995), only a low level of residual hard-seededness
at a similar date, Geraldton had a shorter flowering (< 20%) is needed for persistence of subterranean
period and generally a higher seed yield. Small- clover; the cool climate and summer growth modify
seeded strains of subterranean clover have also been the soil temperature profile, slow the rate of hard-
shown to reach a mature seed weight faster than seed breakdown during summer and enhance the
larger seed lines (Pecetti and Piano, 1994). This survival of seedlings in autumn.
may contribute, at least in part, to the success and Smith et al. (1996) showed that, among legume
dominant position of small-seeded strains of species that have a similar hard-seed level, the pat-
subterranean clover, such as cvs Goulburn, tern of breakdown from hard to soft (permeable)
Denmark and Leura, recent releases from the seed differed, with some breaking down more
Australian breeding programme where high seed rapidly in summer and early autumn, while others
production is considered a high priority. had a delayed pattern. The latter pattern resists false
Rossiter (1966) also identified conservation of breaks, but it may decrease competitive ability with
seed, particularly through hard-seededness, as a rapidly germinating weed species. Thus, in a
crucial trait for success. The importance of hard- summer-dominant rainfall environment, Lodge
seededness has been borne out consistently in (1996) showed that the seedling recruitment of
wheat-belt environments in western Australia subterranean clover and barrel medic was highest
(Smith et al., 1996) and NSW (Wolfe, 1985; Dear when they germinated in midsummer, whereas
et al., 1993a). In western Australia, where the with other legumes it was highest with an autumn
incidence of false breaks (early rains separated from germination.
the main rainfall, resulting in germination of the Other notable characteristics of subterranean
seed and subsequent death of seedlings) is low, clover are those that confer resistance or tolerance
hard-seededness is important to enable subter- to various limiting factors, including diseases, such
ranean clover seed reserves to carry through the as clover scorch (Collins and Stern, 1987) and root
1–2-year cropping phase. In NSW, the cropping rots (Dear et al., 1993b), insect pests (Gramshaw
phase is 4–6 years long and most pastures are et al., 1989), soil nutrient deficiencies/toxicities and
resown, rather than regenerating from hard seed. waterlogging (Reed et al., 1985). These characteris-
However, in NSW, the seasonal ‘break’ is much less tics are important in ensuring the adaptability and
well defined than in the more typical competitive ability of particular cultivars and
Mediterranean climate of south-western Australia strains of subclover in stressful (sensu Grime, 1977)
(Cornish, 1985); false breaks may reduce the num- situations.
Soft seed
soil moisture (0–5 cm)

loss allelopathy
temperature
Newly emerged seedlings

pests and diseases
soil moisture (0–7.5 cm)
light
loss
allelopathy
nutrients
temperature
8–10-week-old seedlings
pests and diseases

high summer loss light
temperatures nutrients
and temperature
temperature
fluctuations
Adult reproductive
plants
pests and diseases

loss light
nutrients
New seed
loss soil moisture (0–60 cm)

light
temperature
Hard seed
Fig. 7.6. Stages in the life cycle of subterranean clover and pathways by which the presence of perennial
pasture plants or their residues may influence the outcome.
As noted by Rossiter (1966, 1977), there are bear on the success or failure of one strain growing
attributes of subterranean clover, other than those with another strain or with other species. For
mentioned above, that are not well understood in example, when the defoliation of a mixture of sub-
terms of their effects on competition between terranean clover strains was infrequent, a later-
strains within a season. Little attention has been maturing, longer-petiole strain (Clare) was able to
given to the vegetative stage properties listed by overtop early-maturing, shorter strains (Seaton
Rossiter (1977); properties such as seedling estab- Park, Daliak) (Hill and Gleeson, 1991). This result
lishment capability, leaf size and petiole length may agreed with an earlier study by Rossiter and Pack
(1972), who found that increased stocking rates establishment. For example, McWilliam et al.
during vegetative growth improved the competitive (1970) found differences between legumes and
performance of cv. Geraldton, a short-stature grasses in the water absorption of seeds, rates of
strain, by restricting the overtopping effect of the germination, and early root elongation; in their
taller Woogenellup cultivar. experience, annual ryegrass was superior to annual
Cocks et al. (1982) found that the high for- legumes in its ability to germinate under moisture
mononetin content of some subterranean clover stress. The evidence for subterranean clover,
strains was an important characteristic associated reviewed by Rossiter (1966), indicated that, once
with their long-term success in mixed populations, germinated, subterranean clover seedlings were
since sheep prefer to eat cultivars with a low content more susceptible to moisture stress than common
of formononetin (Rossiter, 1974). More recently, associate species, such as barley grass, capeweed,
Cocks (1992) found that levels of genistein, another erodium and Paterson’s Curse. According to
of the three phyto-oestrogens in subterranean clover, Rossiter (1966), non-legumes were robust in years
tended to increase with time in divergent genotypes in which ‘false breaks’ occurred in mid-autumn,
of subterranean clover in the field. Genistein is while the contribution of subterranean clover
thought to be one of the phytoalexins, substances declined as a result of insufficient numbers of
produced by plants to activate enzyme defences in surviving plants. However, if the early break was
response to infection by fungi and bacteria (Bell, followed by moist conditions, subterranean clover
1981, reported in Cocks, 1992). was favoured; late rains favoured a range of species
In summary, the ability of subterranean clover including subclover (Rossiter, 1966).
to survive and compete in plant communities is In addition to population density, phosphate
determined by a number of known factors, of supply and defoliation regimes also affect the
which time to reach maturity and hard-seededness botanical composition of annual pastures in southern
are paramount. However, the influence of some Australia (Rossiter, 1966). In southern NSW, a
other characteristics of the species on competitive- series of investigations on ley pastures grazed by
ness is not precisely known. sheep (Ayres et al., 1977) revealed that subter-
ranean clover dominated heavily grazed leys at
COMPETITION BETWEEN SUBTERRANEAN CLOVER AND moderate levels of phosphate supply and low levels
OTHER ANNUAL SPECIES. There have been a number of of soil N, but barley grass quickly became dominant
long-term studies undertaken on the dynamics of if superphosphate was top-dressed at moderate to
annual pastures in the Mediterranean climate of high rates (> 150 kg ha1 annually), particularly if
southern Australia (Rossiter, 1966; Cocks, 1994a). stocking rates were low. This and other work on
These studies are relevant, at least in an ecological defoliation (Collins, 1978) and management
sense if not strictly an agronomic one, to species fre- (FitzGerald, 1976) of subterranean clover led on to
quency and productivity in short-term subterranean recommendations for clover leys in southern NSW
clover leys that regenerate or are sown after a cropping (Southwood and Wolfe, 1978). These recommen-
phase. The conditions at the end of the cropping dations, which are still current (Dear and Sandral,
phase, such as depleted levels of soil N and low levels 1997) were: choose a persistent cultivar of subter-
of disease organisms, favour the early growth of ranean clover; establish at 3–10 kg ha1 under a
clover. Thereafter, the success of subterranean clover light seeding rate (< 20 kg ha1) of a cover crop
during the pasture phase is determined by factors (barley); fertilize the cover crop with additional
such as the size of the populations of other annual superphosphate for the pasture phase (a total of
grasses and herbs, seasonal conditions, phosphate 180 kg ha1, drilled); avoid or minimize top-
supply and defoliation/grazing (Rossiter, 1966). dressed phosphate during the pasture phase by
In southern Australia, the main competitors monitoring the trend in soil phosphate levels; graze
with subterranean clover in annual ley pastures are leys continuously with at least moderate stocking
barley grass, annual ryegrass, silvergrass, brome rates of sheep during the vegetative phase of
grasses (Bromus spp.), capeweed and growth; and avoid excessive defoliation (grazing,
erodium/storksbill (Erodium spp.) (Rossiter, 1966). hay cuts) during spring.
Low annual legume contents in mixed pastures can In recent years, the population balance has
be traced to differences between non-legumes and probably swung towards subterranean clover due to
legumes in a number of characters associated with the availability and use of clover cultivars that are
more persistent than the ones they replace (Dear development of non-chemical methods for control-
and Sandral, 1997) and to the application of herbi- ling vulpia is now a priority, but progress may be
cides to reduce the presence of pasture species that are slow. The findings of Leys et al. (1993) and
weeds during the cropping phase (Powles et al., Dowling et al. (1997) indicated that competition
1997). One species that declined was annual ryegrass, from other species was by itself insufficient to sup-
which, from the 1970s, was the target of a range of press vulpia (Table 7.8). Although increasing the
selective herbicides, but it has now developed density of clover reduced seed set by vulpia in the
resistance to most of these chemicals (Powles et al., first year, there was no difference between the den-
1997). Another species that declined was barley grass, sity treatments by the third year; the inclusion of a
which was a common associate of subterranean clover competitive grass (annual ryegrass, itself a weed)
in well-fertilized pastures (Ayres et al., 1977); lower was the most effective strategy in the experiments
rates of superphosphate applied to leys, higher stock- of Leys et al. (1993). Dowling et al. (1997) con-
ing rates and herbicides were the probable factors that cluded that the key components of an integrated
led to a lower frequency of this grass. control strategy for vulpia were a persistent clover
Conversely, silvergrass (Vulpia spp.) has strain, the inclusion of a competitive grass, forage
apparently increased in frequency in ley pastures, conservation and other seed capture techniques and
presumably due to less competition with ryegrass and grazing management.
higher stocking rates. According to Leys et al. (1991)
and Dowling et al. (1997), the incidence of vulpia has SUBTERRANEAN CLOVER IN MIXTURES WITH PERENNIAL
increased substantially in the last decade, primarily SPECIES. Most studies of the ecology of subterranean
through its tolerance of selective herbicides, which clover have involved monocultures and few have
have removed one of its competitors, annual ryegrass, studied its ability to coexist with perennial species.
and the adoption of reduced tillage practices, which A summary of the stages and processes by which
leave the seed bank intact near the soil surface. Other perennials may have a potential impact on the
factors that have favoured an increasing incidence of annual legume life cycle is presented in Fig. 7.6.
vulpia and attributes that contribute to its competi- There are two main environments where subter-
tiveness are listed in Table 7.7. ranean clover is grown with perennials. The first is
The herbicide simazine is one of the widely in predominantly permanent pastures in the higher
used options to control vulpia but there is concern rainfall (> 500 mm annual rainfall), elevated table-
that the current use of simazine on triazine-tolerant land environments of NSW, Victoria and Tasmania,
canola varieties and on lupin crops and its possible where subterranean clover occurs with phalaris or
use to winter-clean pastures will hasten the devel- cocksfoot. The second is in parts of the cropping
opment of simazine-tolerant vulpia biotypes. The zone where subterranean clover has been sown in
Table 7.7. Factors favouring dominance of Vulpia spp. in pasture swards,

southern NSW (based on Leys et al., 1993; Dowling et al., 1997).
Management and edaphic factors

Reduced use of disc plough
Widespread use of selective herbicides to control annual ryegrass
Reduced vigour of annual legumes due to pathogens, lower fertilizer use
Heavy grazing
Allelopathic effects of plant residues on annual legumes
Transfer of seed in hay
Lower soil pH
Occurrence of droughts
Characteristics of Vulpia contributing to competitiveness
High seed production capacity
Low palatability
Delayed early growth
Multiple germinations, seed dormancy
Table 7.8. Effect of subterranean clover density on Vulpia invasion in pastures at Wagga Wagga, south-
ern NSW (Leys et al., 1993).
Vulpia density and seed production m2
1990 1991
1992
Pasture density/composition Plants Seeds Plants Seeds Plants
Low subclover density 720 1,071,300 22,610 432,000 9,200

Medium subclover density 860 536,200 13,370 399,700 7,700
High subclover density 550 239,300 8,890 386,100 9,720
Medium subclover density + annual ryegrass 640 179,400 4,290 173,300 2,530
combination with lucerne as part of 4–6-year ley perennial plants (Dear, 1998). Seed production by
phase between cropping phases. subterranean clover is known to be sensitive to
The ability of subterranean clover to form stable shading (Collins et al., 1978). In Dear’s (1998)
mixtures in swards with perennial grasses has been experiments, the presence of perennials did not
poor (Dear, 1998). Typically, the annual legume apparently increase the level of moisture stress expe-
content declines over time, leading to grass-domi- rienced by the companion annual legumes in
nant, N-deficient, less productive swards. There are spring.
a number of possible reasons for the inability of An extreme case of the shading effect may
subterranean clover to respond to the decreasing account for the strong competitive advantage of
grass vigour as N availability declines. The hard- white clover over subterranean clover. The combi-
seed breakdown of the subterranean clover seed nation of these two species is most likely to occur
bank may be depressed by the smaller diurnal tem- in environments such as the northern tablelands of
perature fluctuations at the soil surface, due in part NSW, where at least 60% of the total annual rain-
to the increased cover of residual herbage. Another fall (> 700 mm) occurs in summer, outside the
view is that allelopathic chemicals released by the growing period of the annual. The studies of Smith
grass residues inhibit the germination and survival and Crespo (1979) and Hill and Gleeson (1990)
of subterranean clover seedlings (Leigh et al., pointed to the ability of white clover to elongate its
1995). More recently, it has been shown that the petioles in spring and shade the companion subter-
seedling survival and early growth of subterranean ranean clover, resulting in large reductions in seed
clover can be greatly reduced by competition for set of the annual. Another important factor could
moisture with perennials; perennials such as lucerne be the large losses in subterranean clover seed
or phalaris decrease the favourable period of soil reserves that can occur when the environment
moisture following a rainfall event in late summer remains moist after seed set (Archer, 1990).
or early autumn (Dear and Cocks, 1997). This
deficiency leads to a more rapid desiccation and SUMMARY. In subterranean clover pastures, recent
smaller size of clover seedlings and increased work has filled in a number of gaps in our under-
seedling mortality. Furthermore, a combination of standing of competition between and within
shading by the perennials and lower available species. Higher stocking rates, lower rates of super-
nitrate levels in the presence of perennial grass phosphate applied to ley pastures, shorter leys, the
decreased clover seedling growth rates (Dear et al., release of improved cultivars and winter cleaning
1998). are factors that, if anything, are making it easier for
In spring, also, competition between subter- farmers to grow legume-dominant subterranean
ranean clover and perennials (phalaris, lucerne) was clover pastures. However, the advent of herbicide-
found to be important – the quantity of seed set by resistant weeds and the increasing use of perennial
subterranean clover was inversely related to the species are issues that could intensify competition
amount of light intercepted by the canopy of between subterranean clover and non-legumes.
Species dynamics in annual of the shallow burial of seed is the improvement in

Medicago pastures seed–soil contact, which assists in uptake of water
when rainfall is light (Cocks, 1993). Seeds in surface
The available reviews of medic pastures (Cocks et
pods often fail to germinate with light falls of rain
al., 1980; Crawford et al., 1989) place less emphasis
and, although this can be an advantage in that it
on the competitive ability of medics than on the
prevents premature germination and the depletion
ability of individual species to grow, produce seed
of the seed bank, it can shorten the growing season
and persist. This emphasis is understandable, since
and reduce dry-matter yield in good seasons
in Australia medics are traditionally grown in short-
(Cocks, 1993).
term pasture/crop leys, where the survival and
The benefit of sowing mixtures of annual
regeneration of the medic after the cropping phase medics in Australian farming systems has not been
is of paramount importance (Puckridge and widely reported. Latta and Carter (1996) con-
French, 1983). Another possible constraint to long- cluded that there were benefits in combining an
term studies of species relationships in medic early-maturing small-seeded cultivar, such as
pastures is the medic decline syndrome (see Table Harbinger AR, with a mid-season, larger, soft-
7.2), which was first reported in the 1980s (Carter seeded cultivar (Paraggio), to maximize seed
et al., 1982) after a devastating occurrence of the production and regeneration. The relationships
spotted alfalfa aphid and the blue-green aphid in between annual medics and competing species
the late 1970s. This decline has continued in should be included in any future monitoring
pastures in South Australia, where low soil phos- programme to evaluate the long-term contribution
phorus and zinc levels and Pratylenchus nematodes of these newer varieties to agriculture.
have also been implicated (Denton and Bellotti,
1996). The extensive use of residual sulphonylurea
herbicides is another factor considered to affect the Increasing the diversity and stability of
persistence of annual Medicago spp. grown in annual pasture swards
rotation with crops. In recent years, medic breeding
and evaluation in Australia have been dominated by The legume improvement programmes in Australia
the need to develop aphid-tolerant varieties; the are changing their emphasis after 50 years of
programme has resulted in the release of new barrel mainly concentrating on developing new cultivars
medics (M. truncatula), cvs Parraggio, Sephi, of subterranean clover and annual medics. They are
Caliph and Mogul, and strand medics (Medicago being refocused towards new, previously unculti-
littoralis), cvs Harbinger AR and Herald. vated, annual legume species. This development in
The vulnerability of medic pods to ingestion by part reflects the significant progress that has been
sheep and goats is a critical factor determining the made in subterranean clover towards the objectives
amount of the germinable seed pool present in of lowering oestrogens, increasing hard-seededness
autumn–winter, thereby influencing the ability of and improving resistance to leaf and root diseases
the medic to maintain high plant populations in by breeding programmes (Nichols et al., 1994). But
longer term pastures. Dear and Jenkins (1992) it also reflects a need to find species adapted to
found that sheep could remove 200–600 kg ha1 niches not suited to subterranean clover and to
medic seed in 7 days when grazed at a high stock- introduce new, adaptive mechanisms not contained
ing rate. The greater proportion (up to 95%) of the within subterranean clover or annual Medicago spp.
seed is digested, with the recovery of seed in sheep These include greater adaptation to waterlogging,
pellets being inversely proportional to the size of different patterns of hard-seed breakdown, small
seed (Russi et al., 1992). seeds that can pass through the rumen undigested,
Shallow seed burial has been shown to enhance aerial seeding for ease of harvesting and greater
the rate of softening in the first year (Cocks, 1993), insect tolerance. A good example of how other
as well as preventing ingestion of the pod by stock. species may fill gaps in pastures is the colonization
Hence most successful farming systems involving by T. clusii Godr. & Gren. of poorly drained niches
medics require frequent cropping and shallow seed within subterranean clover pastures (Cocks,
incorporation. Deep ploughing, as commonly prac- 1994a). Cocks (1994a) has suggested that the lack
tised in western Asia and North Africa, results in of species diversity in Australian pastures may be
poor regeneration (Cocks, 1994b). Another benefit slowly resolving itself through a combination of
hybridization, mutation and the spread of species implications for breeding and selection pro-
by grazing animals. This process can be accelerated grammes and how new species are evaluated.
by carefully targeting selection and introduction Evaluating species in monocultures in small plots
programmes. may be inappropriate, and it may be preferable to
Equally importantly, the change in emphasis is evaluate combinations of species under realistic
also an attempt to increase the diversity of species grazing regimes to discover their dynamics, produc-
in pasture swards. In response to temporal and tivity, stability and resilience. Such an approach is
spatial variation, diversity may produce small but in line with proposals by Cocks (1992), who con-
important improvements in ecosystem stability and cluded that, due to the complexity of the possible
resilience, at least in terms of biomass, if not in interactions, the evaluation of species mixtures
botanical composition (Tilman, 1996). Diversity is should be conducted in farmers paddocks under
also a step away from the annual legume monocul- practical management conditions.
tures, which have dominated Australian pastoral
systems, monocultures that have been vulnerable to
diseases, such as clover scorch (Kabatiella caulivora) Conclusions
or root rots in subterranean clover, or pests, such as
the spotted alfalfa aphid. This review has highlighted the main developments
The use of more diverse mixtures of legume that have occurred and how they have shaped the
species opens up a whole new area of plant compe- dynamics of species relationships in southern
tition, which has received very limited study. While Australian pastures. The main phases were a century
many of the factors identified earlier in competition and a half (1800–1950) of exploitation of Australian
between genotypes of the same species may be native grasslands (a phase that is continuing, albeit at
important, other factors will be involved. For example, a slower rate), a short and intensive period
a study by Dear and Coombes (1992) found that (1950–1970) of pasture improvement with fertilizers
when T. subterraneum, Medicago murex and and exotic species and a period of re-evaluation
Trifolium michelianum Savi were grown in binary during the last 30 years of the 20th century. This last
mixtures, seed production by T. michelianum was, period has provided an opportunity to consolidate
relative to the monoculture yields, more sensitive to the knowledge of scientists and the experience of
competition than either of the other species, setting farmer/graziers and to reflect on successful and
only 59–81 kg ha1 in mixtures, compared with unsuccessful aspects of pasture management in
287–514 kg ha1 by the other two species. relation to the future of permanent and ley pastures
Whether this was due to the crash grazing system in Australian agriculture.
employed or the shading of one species by another In this review, a number of gaps in knowledge
could not be determined. Another finding was the were indicated. First, there was a lack of knowledge
need for heavy grazing over summer for T. miche- on the effects of ‘management’ (subdivision, grazing
lianum to regenerate satisfactorily the following and fertilization) on plant communities, covering a
autumn (Dear and Coombes, 1992). In contrast, representative range of native, naturalized and
the seed reserves of larger-seeded species, such as exotic species. Fortunately, work is under way to
M. murex, contained in a large pod on the soil enhance the understanding of agriculturists,
surface, can be severely depleted through consump- ecologists and pastoralists on the separate effects of
tion by livestock over summer, resulting in poor defoliation, trampling and nutrient addition/deple-
recruitment the following year. tion on plant–soil–water relations in native and
These findings reinforce the challenge of choosing improved grasslands. A feature of this work is a
a grazing strategy that maintains the diversity, prof- greater emphasis on the impact of species on soil
itability and sustainability of pasture mixtures. In erosion and on the hydrology of catchments
extensive areas where grazing management is less (Johnston et al., 1999).
well controlled, it may be more important to select Secondly, there is a need for a more theoretical
species combinations that can survive under a approach to grassland dynamics, in which the
particular grazing regime rather than attempting to observed changes in productivity and botanical
force species together with divergent grazing needs, composition are related to useful frameworks that
despite the desirability of other agronomic charac- might help integrate knowledge and explain
teristics. These combinations have important phenomena. In this respect, the application of the
state and transition model (Humphreys, 1997) to pasture types, such as those based on annual
plant succession in permanent pastures and of the medics, must be overcome.
CSD model (Grime, 1977) to ley pastures is rec- In Australian croplands, developments such as
ommended. Both models eschew the dogma that is the changing spectrum of crop weeds and the
associated with the traditional Clementsian (linear) advent of herbicide resistance are likely to force
model of succession, and they are flexible enough agronomists and managers to place a greater
to encompass not only breadth of knowledge but value on monitoring, understanding and manip-
also the targeting of a particular problem or issue. ulating the population dynamics of pasture/weed
Thirdly, the review has pointed to a lack of species, during both the cropping and pasture
quantitative evidence available for the decline in phases of rotations. In grazing lands, too, the
legume and total pasture productivity that has scene is changing, due to enhanced community
reportedly (Carter et al., 1982) occurred in pastures perceptions of the sustainability of catchments
in southern Australia. A recent survey (Australian and the aesthetic value of landscapes.
Bureau of Agricultural and Resource Economics Developments such as new pests and diseases or
(ABARE), 1999, personal communication) revealed the loss of key herbicides due to community or
that farmers in the cropping zone of Australia were corporate imperatives will ensure the need for
predominantly satisfied with both the productivity theoretical and applied studies of plant competi-
and quality of their pastures. The separate views of tion and pasture dynamics.
researchers and farmer/graziers need further explo-
ration and, as suggested by Wilson and Simpson
(1993), there is scope for regular and standardized Note
surveys of the state of pastures. Strategic long-term
studies of pasture relationships, similar to those 1. In Australia, there is a trend towards the use of
undertaken by Hutchinson and colleagues ‘ley’ to denote a 1-year pasture and ‘phase’ to
(Hutchinson, 1992; Hutchinson et al., 1995; denote several years of pasture, but we prefer the
Hutchinson and King, 1999) on permanent pas- use of the terms ‘ley’ and ‘ley farming’ to embrace
tures on the northern tablelands of NSW, are pastures of one to several years’ duration in the
warranted. The lack of such studies for some crop/pasture rotation.
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C&S in Pastures Chap 07 22/10/01 1:38 pm Page 119

7 The Population Dynamics of Pastures,

E.C. Wolfe1 and B.S. Dear2

Introduction the dynamics of plant communities. Then, in a

© CAB International 2001. Competition and Succession in Pastures

120 E.C. Wolfe and B.S. Dear

Table 7.1. Agricultural development in Australia, 1820–2000.a

Livestock numbers ( 106)

Exploration 1820 0.3 < 0.1 0.01

Population Dynamics of Pastures 121

122 E.C. Wolfe and B.S. Dear

Population Dynamics of Pastures 123

Original climax community

more intense grazing

Dwarf cool-season perennials + dwarf warm-season perennials

Stipa falcata, Chloris truncata

Dwarf cool-season perennials + cool-season Mediterranean annuals

phosphorus addition and grazing

Current pasture ley community with sown species,

124 E.C. Wolfe and B.S. Dear

Population Dynamics of Pastures 125

Medic decline, South Australia (Carter et al., 1982)

E.C. Wolfe and B.S. Dear

–F Improved +F Degraded by acidity,

Population Dynamics of Pastures 127

128 E.C. Wolfe and B.S. Dear

Population Dynamics of Pastures 129

130 E.C. Wolfe and B.S. Dear

Population Dynamics of Pastures 131

Pasture species Area (million ha) % of freehold + leasehold land

Table 7.5. The relative importance (on an area basis) of pas-

Description Percentage by area

Unimproved native pasture 62.5

132 E.C. Wolfe and B.S. Dear

LIMITS TO ADAPTATION (TEMPERATE SPECIES)

a persistent perennial grass component in pasture Phalaris – a persistent grass in Australia

Population Dynamics of Pastures 133

Low stocking Medium stocking High stocking

Phalaris – basal cover (%)

Phalaris – basal cover (%)

134 E.C. Wolfe and B.S. Dear

Soil treatment in pot Binary mixtures

phalaris (cv. Sirosa): ED Al50 value = 28 M, sensitive

Population Dynamics of Pastures 135

136 E.C. Wolfe and B.S. Dear

Population Dynamics of Pastures 137

soil moisture (0–5 cm)

Newly emerged seedlings

pests and diseases

pests and diseases

loss soil moisture (0–60 cm)

138 E.C. Wolfe and B.S. Dear

Population Dynamics of Pastures 139

Table 7.7. Factors favouring dominance of Vulpia spp. in pasture swards,

Management and edaphic factors

140 E.C. Wolfe and B.S. Dear

Vulpia density and seed production m2

Low subclover density 720 1,071,300 22,610 432,000 9,200