Agricultural Systems 194 (2021) 103272

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Agricultural Systems
journal homepage: www.elsevier.com/locate/agsy

Selecting higher nutritive value annual pasture legumes increases the

profitability of sheep production
Dean T. Thomas a, *, Bonnie M. Flohr b, Marta Monjardino b, Angelo Loi c, Rick S. Llewellyn b,
Roger A. Lawes a, Hayley C. Norman a
a
CSIRO Agriculture and Food, Private Bag 5, Wembley, WA 6913, Australia
b
CSIRO Agriculture and Food, Glen Osmond, SA 5064, Australia
c
Department of Primary Industries and Regional Development, South Perth, WA 6156, Australia

H I G H L I G H T S G R A P H I C A L A B S T R A C T

• The nutritive value of annual pasture

legumes varies widely among species
and varieties.
• We calibrated Dry Matter Digestibility
in six annual legume models to repre­
sent their year-round nutritive value.
• Mean Dry Matter Digestibility of pas­
tures ranged from 62% to 73% over the
year among the six species modelled.
• The supplementary feeding cost in a
sheep enterprise decreased by up to 20%
when higher quality legumes species
were grown.
• Differences in nutritive value among
pasture legumes are important in
extensive livestock production.

A R T I C L E I N F O A B S T R A C T

Editor: Dr. Jagadish Timsina CONTEXT: The nutritive value of annual pasture legumes changes with phenology during each season, with a
progressive decrease in feeding value as plants accumulate structural carbohydrates and nutrients are diluted.
Keywords: However, these changes differ widely among annual legume species, affecting the pattern of nutrient availability
Pasture legumes and production in a grazing enterprise.
Nutritive value
OBJECTIVE: This paper set out to examine the implications of differences in the nutritive value of annual pasture
Feeding value
legumes in relation to their economic value for mixed farming systems of southern Australia.
Quality
Sheep METHODS: Calibrated GrassGro™ plant models with data generated from field experiments to represent Dry
Matter Digestibility (DMD) profiles of six annual legume species; Medicago truncatula Gaertn. (barrel medic),
B. pelecinus L. (biserrula), Trifolium spumosum L. (bladder clover), Ornithopus sativus Brot. (French serradella),
Medicago littoralis Rhode ex Loisel. (strand medic) and Trifolium subterraneum L. (subterranean clover). The plant
models were used in a modelling study to support a self-replacing Merino sheep enterprise, which was simulated
at seven locations across southern Australia, where annual pasture species are heavily relied on for livestock
production.
RESULTS AND CONCLUSIONS: Modelled values of DMD of green material, averaged across all phenological
stages, ranged from approximately 62% (French serradella) to 73% (bladder clover), with little variation due to

* Corresponding author.
E-mail address: dean.thomas@csiro.au (D.T. Thomas).

https://doi.org/10.1016/j.agsy.2021.103272
Received 1 March 2021; Received in revised form 1 September 2021; Accepted 2 September 2021
Available online 20 September 2021
0308-521X/© 2021 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/by-nc-nd/4.0/).
D.T. Thomas et al. Agricultural Systems 194 (2021) 103272

location. Pastures with higher nutritive value had a reduced requirement for supplementary feeding of sheep.
Variation in nutritive value had a greater effect on supplementary feeding costs and gross margins than differ­
ences in biomass production among the annual legume species. Differences in the levels of supplementary
feeding required in systems with the different legumes was most pronounced during the period from May to July
(late autumn to winter). For sheep enterprises compared at the same stocking rate, the total supplementary feed
required was 18% lower with bladder clover pastures, and 9% lower in subterranean clover pastures, compared
with an average of the remaining four pasture species.
SIGNIFICANCE: This is the first study to compare the effects of variation in nutritive value among the annual
pasture legumes developed for the soils and climatic conditions of southern Australia. Our results demonstrated
that the magnitude of differences in nutritive value among annual pasture legumes are economically important
and that the establishment of higher nutritive value legumes will reduce supplementary feeding costs in
extensive grazing systems, in some cases even where their biomass production is lower.

1. Introduction
A diverse feedbase is used in extensive livestock enterprises in the
mixed crop and livestock production region of southern Australia, which
has a predominantly Mediterranean-type climate. This is necessary to
provide the best chance of meeting the year-round feed requirements of
animals grown in a seasonal and variable climate, with lower reliance on
supplementary feeding and less risk (Bell et al., 2018; Masters et al.,
2007). Traditionally, annual pastures provide most of the feed in these
systems and are essential to sustain high livestock productivity at a time
of year when energy demand is greatest for lactation and growing young
animals (Behrendt et al., 2011; Moore et al., 2009). Annual pasture le­
gumes have been introduced in these systems to improve the nutritional
quality of forage, fix atmospheric N and contribute to increased soil
fertility by maintaining soil organic matter and improving soil physical
conditions (Howieson et al., 2000; Nichols et al., 2012; Porqueddu et al.,
2016).
The amount, timing and quality of forage produced by pastures, and
otherwise supplied by the seasonal feedbase (e.g. stubbles, shrubs,
forage crops), drives growth, reproduction rates, lifetime productivity
and economic value in livestock systems (Behrendt et al., 2011; Mon­
jardino et al., 2014). However, much of the work in the development of
annual pasture legumes has focused on selecting for the ability to
establish a persistent seed bank and grow reliably and productively in
mixed farming systems (Nichols et al., 2012). Less attention has been
given to comparative analysis of new annual legume species in relation
to their contribution to the feedbase, although there has been research
demonstrating the value of improving aspects of this, such as biomass
production and the quality of pastures, at various phenological stages
(Donnelly et al., 1994; Thomas et al., 2010a). A meta-analysis by Sol­
lenberger and Vanzant (2011) concluded that pasture nutritive value
sets an upper limit for livestock production, whereas the biomass
available drives how close to the limit of livestock production will be
reached. The nutritive value profiles of annual pasture legumes vary
widely among species and across the growing season. For example, the
metabolizable energy content of 15 annual pasture legume genotypes
(whole plant) for a single site ranged from 11.1–13.5 MJ ME/kg when
vegetative, 10.2–12.8 MJ ME/kg when flowering and 8.4–11.0 MJ ME/
kg at senescence (Thomas et al., 2010a). Annual plant species demon­
strate a decline in Dry Matter Digestibility (DMD) as plants mature,
which is presumably associated with lower cell wall content of young
compared with older leaves and stems (Mcleod et al. 1990). Annual
pasture legumes also vary widely in their rate of maturity, with flow­
ering ranging from less than 80 to more than 150 days from sowing
(Nichols et al., 2007). A change in the rate of decline in DMD during
maturity may be as important as the DMD at peak biomass in spring.
However, such pasture characteristics tend to be overlooked in pasture
legume domestication, where new cultivars are developed and released
based on their ability for fill a priority role within a farming system
(Ewing, 2017).
Considering the diverse range of events and interactions in extensive
livestock production to evaluate a systems intervention is a complex
problem. Process models are increasingly used in the design and opti­
misation of livestock systems in order to meet the increasing demands of
the supply chain and consumers for high quality products that are pro­
duced on extensive grazing land in an acceptable manner (Gouttenoire
et al., 2011). In this paper, we used biophysical modelling to compare
the systems outcomes of differing nutritive value profiles for annual
pasture legumes, based on the characteristics of species that are
currently being developed for commercial release. We hypothesised that
the differences in nutritive value among annual pasture legumes would
be a more significant economic driver than differences in biomass
production.
2. Materials and methods
A simulation modelling experiment was conducted to determine the
economic value to a livestock enterprise associated with the energy
density of the dry matter produced in annual pasture legumes. To do
this, we sourced data from field experiments (including unpublished
data) of the nutritive value of annual pasture legumes (Norman et al.,
2020). Values of Dry Matter Digestibility (DMD) and Nitrogen (N) were
aggregated at the species level and any differences in accessions were
ignored. We then fitted DMD profiles for six annual pasture legumes
using existing GrassGro™ plant models that could represent each of the
legumes, calibrating the Beta plant model to minimise the root mean
square error (RMSE) of the fitted line. A basic Beta model for each plant
species was considered sufficient for the purpose of the study, which
aimed to look broadly at how seasonal DMD profiles affect livestock
production and not to evaluate the skill of the Beta models to predict
outside the calibration data. The N content of the pastures was not
adjusted in the Beta plant models. Due to the relatively high N content of
annual legumes at any given DMD, the N requirements of both adult and
young sheep (e.g. 16–20 g N/ kg DM intake) was sufficient to meet the
animal’s requirements (CSIRO, 2007). The plant models were then
implemented in simulation modelling scenarios of a self-replacing Me­
rino enterprise to establish effects on sheep feeding at seven locations
across southern Australia.
2.1. Scenario locations
The seven locations used for simulation of livestock enterprises were
in the low-medium rainfall area of the mixed farming region of southern
Australia, which is the target area for adoption of the new pasture le­
gumes (Fig. 1). Annual rainfall (1989–2018) for the sites was as follows
(mean ± SD); Cascade 402 ± 86 mm, Condobolin 483 ± 142 mm,
Corrigin 338 ± 91 mm, Karoonda 351 ± 102 mm, Lameroo 354 ± 95
mm, Mingenew 337 ± 83 mm, Waikerie 277 ± 96 mm. In these regions,
rotations of crops (cereals, pulses and canola) and pastures are grown,
with Merino sheep grown for wool and meat being the main form of
livestock production. Longer term trends in production in the mixed
farming region are described by Kirkegaard et al. (2011).

2
D.T. Thomas et al. Agricultural Systems 194 (2021) 103272

Fig. 1. Experimental locations in three states of southern Australia used in simulation modelling scenarios.

2.2. Livestock enterprise model chosen time interval (30 years).

A simulation experiment was conducted with GrassGro™ (version
3.3.10; Moore et al., 2007) using six plant models (annual pasture le­
gumes; Table 1) × seven sites (Fig. 1). The GrassGro™ model is a
ruminant grazing model that has been developed for extensive livestock
systems of southern Australia (Moore et al., 1997; Mokany et al., 2010).
GrassGro™ is comprised of components that describe the biophysical
(climate, soils and land management units (paddocks), pastures, live­
stock), managerial (e.g., stocking rate, soil fertility, pasture grazing ro­
tations and animal reproductive management) and financial subsystems,
which form the ‘farm system’ under consideration. The financial anal­
ysis was based on a gross margin analysis, with variable costs and in­
come taken from the Australian Bureau of Statistics over the 5 years,
2016–2020. These components combine to simulate biophysical and
economic performance within the farm system at daily time steps for the
The plant models were calibrated using field data as described in the
next section. A single enterprise type (self-replacing Merino ewes) was
selected for the study. The soil type selected within GrassGro™ was kept
the same for all locations in the scenarios (uniform, coarse textured,
sandy soil: ref. Uc5.22; Northcote, 1979), and pastures were grown as
monocultures of the tested annual legumes. In the GrassGro™ model,
different legume species are not sensitive to differences in soil texture as
is seen in practice. In the model, sheep were rotated between annual
green and dry legume pastures, stubbles (December and January) and
shrubs (March) during each season. Feed from crop stubbles and
perennial saltbush pastures was kept constant and the deficit (not met by
annual pasture) was made up by supplementary feeding, which varied
by season, location and pasture species. Therefore, the higher quality
pastures were able to displace supplementary feeding both by extending
the period where edible biomass is available and allowing sheep to gain

Table 1
Agronomic characteristics of the legume species that were used in this study.
Annual pasture legume species Soil nichea Example cultivar, release (year) Sowing to flowering (days) Hard seed (%)

Barrel medic Heavier alkaline soils Paraggio, 1982 98–110 65–75

Subterranean clover Sandier, more acidic soils Dalkeith, 1983 97 10–20
Biserrula Fine textured acidic soils Casbah, 1997 100–105 90–95
French serradella Deep sands with poor water holding capacity Cadiz, 1997 90–115 0–10
Strand medic Heavier alkaline soils Angel, 2005 70–90 85–95
Bladder clover Fine textured acidic and alkaline soils Bartolo, 2008 105–110 60–70
a
Adapted from Nichols et al., 2007 and informed by local pasture agronomists.

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D.T. Thomas et al. Agricultural Systems 194 (2021) 103272

Table 2
Overview of the experimental sites (species × year of establishment) from which data was sourced for use in calibrating plant models.
Year Barrel medic Biserrula Bladder clover Serradella Strand medic Subterranean clover

2012 3 3 3 0 0 6
2013 3 4 6 1 0 0
2014 0 1 0 1 0 0
2018 0 0 0 8 8 0

condition more rapidly so that they took longer to reach condition
thresholds where supplementary feeding was required. A single stocking
rate (7 breeding ewes per winter grazed hectare) was selected across
sites, based on results of an optimisation function for stocking used in
previous research (Thomas et al., 2019). All simulations were run over
30 years (1989–2018) using historical weather data from the Australian
Bureau of Meteorology.
white clover model behaved as a long-season, semi- or indeterminant
annual pasture species due to seasonal terminal drought conditions. The
sowing dates of field experiments differed according to seasonal con­
ditions and experimental design, so the number of days from sowing was
used to standardise across the experiments. Therefore, differences in
plant phenology associated with sowing date were not fully accounted
for in the plant models.

2.3. Plant model calibration 3. Results

The GrassGro™ simulation model contains a library of para­
meterised plant models, that represent the major forage species of
southern Australia. Three of the existing GrassGro™ plant models were
used as base models to calibrate Beta models that represented the six
annual pasture legumes evaluated, using Dry Matter Digestibility data
that was sourced as described earlier in the methods. Calibration of each
of the plant models was conducted using experimental data from field
sites in the mixed farming regions of Western Australia and South
Australia, collected in four years between 2012 and 2018 (Table 2).
Existing GrassGro™ base models were assigned to the Beta models as
follows: i) subterranean clover cv. Dalkeith, for subterranean clover
(Beta) and bladder clover (Beta); ii) barrel medic cv. Paraggio, for barrel
medic (Beta) and strand medic (Beta); and iii) white clover for biserrula
(Beta) and French serradella (Beta v2). French serradella (Beta v1) had
been previously calibrated with field data of plant growth and used the
same parameter values, except DMD, as reported in Table 3 and Fig. 2.
Calibration data were assigned to four ‘days from sowing’ categories
(Table 4) and DMD values for k-q-leaf and k-q-stem parameters
(described in Table 3) were adjusted in the each of the plant models.
Indicative results of model calibration for bladder clover (Beta) and
French serradella (Beta v2) plant models are reported in Table 4.
Although white clover is developed as a perennial plant model in
GrassGro™, plant reproduction is also based on a seed biomass pool. As
a result, under the climatic conditions of the GrassGro™ scenarios the
3.1. Plant model testing
Leaf and stem quality parameters selected for the base and novel
annual legume plant models based on calibrating the model using
measured field data are presented in Table 5. The main differences in
parameter values were minimum DMD values of leaf but particularly
stem at the reproductive stage of maturity (kq-leaf 3 and kq-stem 3),
which highlights the high variability in the rate of decline in DMD
among species.
The RMSE for modelling values for the fitting of the simulation data
are presented in Table 5, with fitted plots of the field data and calibrated
lines of modelled DMD across the season (days from sowing) displayed
in Fig. 3. Overall, the models represented the change in DMD of the
above-ground pasture (shoot) biomass well. In the case of biserrula and
bladder clover, there were substantive differences in the seasonal
nutritive value profiles among field studies and the model calibration
tended to fit the field data with the lower DMD profiles. It was difficult to
calibrate the model to the very low nutritive value of senesced material
that was observed for the medics and subterranean clover (Fig. 3).
The plant models were not calibrated for protein content. Testing of
crude protein (N × 6.25) output from the models using data published
by Thomas et al. (2010a) indicated the existing nitrogen (N) values used
in the plant models represented observed values variably among the
pasture species (Table 6). Modelled N values for barrel medic and French

Table 3
A summary of plant model parameters used in French serradella v2.0 with parameter descriptions.
Code Assigned value Parameter description

k-v-3 0.0 Base temperature for degree-day computations (◦ C)

k-v-5 no value Degree day sum for commencement of reproductive growth (◦ d)
k-v-6 700 Degree day sum for commencement of flowering (◦ d)
k-v-20 3.0 Length of the drought period required to induce senescence (d)
k-v-21 3400 Value of degree days to senescence in the absence of drought (◦ d)
k-i-1 0.035 Reference specific leaf area (ratio of LAI to leaf weight) (m2/g)
k-i-7 0.7 Apparent light extinction coefficient (ungrazed conditions) (0–1)
k-i-8 0.8 Apparent light extinction coefficient (grazed conditions) (0–1)
k-tl-3 0.05 Maximum proportion stem mass at flowering to be relocated to seed (0–1)
k-r-1 1500 Maximum rooting depth under optimal soil conditions (mm)
k-r-2 2.0 Maximum rate of root front extension (mm/◦ d)
k-s-1 0.06 Rate of “hardening” of immature seeds (/d)
k-s-2 100 Length of period of innate dormancy (d)
k-g-6 8 Time to first seedling emergence under optimal conditions (d)
k-g-7 15 Time to complete seedling emergence under optimal conditions (d)
k-q-leaf-1 0.8 Average DMD of newly produced leaf (g/g)
k-q-leaf-2 0.7 Minimum DMD of green leaf during vegetative growth (g/g)
k-q-leaf-3 0.7 Minimum DMD of green leaf during reproductive growth (g/g)
k-q-stem-1 0.8 Average DMD of newly produced stem (g/g)
k-q-stem-2 0.6 Minimum DMD of green stem during vegetative growth (g/g)
k-q-stem-3 0.4 Minimum DMD of green stem during reproductive growth (g/g)

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D.T. Thomas et al. Agricultural Systems 194 (2021) 103272

3000 NPP among the pastures modelled was similar within sites except for
biserrula and French serradella, which had 15 and 18% higher NPP on
average across the sites compared with the other pastures. The differ­
2500
ences between French serradella and biserrula compared with the other
species were greatest at the Cascade and Condobolin sites, and smaller at
2000 Corrigin and Mingenew, which have shorter growing seasons (Fig. 4).
The DMD of green (across the growing season) and dry (immediately
Shoot
Biomass post-senescence) total above-ground plant material for each of the
1500
(kg/ha) annual pasture legumes, averaged across all sites, is reported in Fig. 5.
There were clear differences in DMD within sites. Bladder and subter­
1000 ranean clover had higher DMD, while barrel medic and French serra­
della had the lowest average DMD, corresponding with the plant model
calibration (Table 4). The DMD of bladder clover immediately post-
500 senescence was higher than the average of live French serradella
(Fig. 5). While there were clear differences in DMD among the pasture
0 legumes within sites, differences were relatively small between sites.
Jun Jul Aug Sep Oct Oct Nov Dec Sites with a longer growing season, such as Condobolin and Cascade,
tended to have higher average DMD in living material than sites with a
Fig. 2. Observed (■, error bars are standard errors of the means) Ornithopus short growing season (e.g. Mingenew).
sativus cv. Cadiz from Thomas et al. (2010a) and simulated (French serradella The simulated average annual metabolisable energy intake of the
Beta v1) above-ground biomass during 2000 at Wundowie, Western Australia.
sheep at each site is shown in Fig. 6, separated into the 4 feedbase
components modelled; legume pasture, stubble, shrub and supplement
serradella were comparable to the observed data, but N content of feed. Excluding Waikerie, the proportion of energy intake from pasture
bladder clover and subterranean clover was higher than observed legumes ranged from 50% (French serradella at Mingenew) to 68%
values. (bladder clover at Condobolin). The proportion of energy intake from
pasture legumes at Waikerie was much lower (35–42%), with a corre­
3.2. Location × species analysis sponding higher level of supplementary feeding required (about 40% of
Metabolisable Energy intake). Bladder clover pastures made up a higher
The average annual net primary production (NPP, kg/ha), being the proportion of the feedbase across all sites, and within sites were up to
total above-ground biomass accumulated minus losses to metabolism 10% units higher in their proportion of the feedbase represented as
and maintenance, of the six species produced at each location is pre­ Metabolisable Energy intake (Fig. 6). The energy intake of the sheep per
sented in Fig. 4. Overall, NPP corresponded with the total growing unit of biomass production varied widely among the legumes, reflecting
season rainfall and the length of growing season for these locations. The

Table 4
Calibration results for bladder clover (Beta) and French serradella (Beta v2) for Dry Matter Digestibility (DMD) aggregated in four groups of days from sowing, where
kq-stem 1, 2 and 3 values were adjusted (column headings).
Days from sowing Measured DMD% Modelled DMD (g/g) (kq-stem1,2,3a)

Bladder clover (Beta) 0.8, 0.7, 0.7 0.8, 0.7, 0.6 0.8, 0.75, 0.6
80–109 74.1 75.3 73.6 74.5
110–139 72.3 73.2 70.3 71.6
140–169 69.8 70.6 66.9 67.9
>170 65.3 68.4 64.7 64.9
French serradella (Beta v2) 0.75, 0.6, 0.5 0.75, 0.6, 0.45 0.7, 0.5, 0.5 0.7, 0.6, 0.4
80–109 65.1 69.5 67.6 64.6 66.0
59.6
110–139 59.5 64.2 60.8 59.6 61.4
140–169 55.6 59.0 54.6 55.1 56.2
>170 42.5 57.9 53.1 54.0 49.8
a
kq-stem 1 is the average DMD of newly produced stem, kq-stem 2, 3 is the minimum DMD of green stem during vegetative and reproductive growth, respectively.

Table 5
Parameters used in the GrassGro™ plant model for leaf and stem quality (Dry Matter Digestibility, 0–1), and root mean square error (RMSE) of fitted field data points.
Plant model kq-leaf1 kq-leaf2 kq-leaf3 kq-stem1 kq-stem2 kq-stem3 RMSE
(%DMD)

Base models
Barrel medic cv. Paraggio 0.8 0.7 0.7 0.8 0.7 0.6
Subterranean clover cv. Dalkeith 0.8 0.7 0.7 0.8 0.7 0.6
White clover (cultivar unspecified) 0.8 0.7 0.7 0.8 0.7 0.6
Beta models
Barrel medic (Beta) 0.8 0.7 0.6 0.8 0.5 0.4 6.18
Biserrula (Beta) 0.8 0.7 0.7 0.7 0.7 0.5 6.37
Bladder (Beta) 0.8 0.75 0.7 0.8 0.75 0.6 3.83
French serradella (Betav2.0) 0.8 0.7 0.7 0.7 0.6 0.4 5.15
Strand medic (Beta) 0.8 0.8 0.7 0.8 0.7 0.35 6.92
Subterranean clover (Beta) 0.8 0.8 0.7 0.8 0.7 0.5 6.31

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D.T. Thomas et al. Agricultural Systems 194 (2021) 103272

Fig. 3. Simulated (line) and measured (symbol) Dry Matter Digestibility (%) across the season (days from sowing) of annual legumes a) barrel medic b) biserrula c)
bladder clover d) French serradella e) strand medic f) subterranean clover. Within graphs, symbols identify different trial plots where measured data was sourced,
labelled (symbol, plot reference, year) as follows: a. □ SA69 2013, ○ SA69 2012, ∆ SA7 2013, + SA7 2012, × SA74 2013, ◊ SA74 2012 (Urrbrae, South Australia) b.
□ WA BrooktonA 2013, ○ WA Brookton B 2014 (Brookton, Western Australia); ∆ SA10 2013, + SA147 2012, × SA18 2012, ◊ SA20 2013, ∇ SA20 2013, SA47
2012 (Urrbrae, South Australia); c. □ SA11 2013, ○ SA176 2013, ∆ SA176 2012, + SA29 2013, × SA48 2013, ◊ SA79 2013, ∇ SA79 2012, SA93 2012, * SA93
2012 (Urrbrae, South Australia); d. □ WA BrooktonA 2013, ○ WA BrooktonB 2014 (Brookton, Western Australia), ∆ SA13_1 2018, + SA20_2 2018, × SA30_2 2018, ◊
SA41_3 2018, ∇ SA43_3 2018, SA54_4 2018, * SA58_4 2018, ◆ SA7_1 2018 (Urrbrae, South Australia); e. □ SA10_1 2018, ○ SA19_2 2018, ∆ SA2_1 2018, +
SA25_2 2018, × SA40_3 2018, ◊ SA44_3 2018, ∇ SA47_4 2018, SA48_4 2018 (Urrbrae, South Australia); f. □ SA143 2012, ○ SA166 2012, ∆ SA43 2012, + SA45
2012, × SA72 2012, ◊ SA76 2012 (Urrbrae, South Australia).

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D.T. Thomas et al. Agricultural Systems 194 (2021) 103272

Table 6 higher quality pastures to livestock production, such as higher ewe

A comparison of observed and modelled crude protein content (N × 6.25, %) of
green (live) and senesced (dry) pasture legumes.
Barrel medic Bladder clover Serradella Subterranean clover
Green
Observed 21.9 16.5 19.9 17.9
Modelled 19.8 26.1 18.3 23.3
Senesced
Observed 8.0 14.4 11.1 11.0
Modelled 10.1 20.1 9.3 14.3
differences in the DMD profiles of the plants. For French serradella it was
less than 3 MJ ME intake/kg NPP, compared with almost 4.5 MJ ME
intake/kg NPP for bladder clover. Differences in the energy intake of
sheep per unit of biomass produced between sites were relatively small.
In Table 7, average daily pasture intake is separated into the DMD
pools (30–80% DMD) that are used within the GrassGro™ model to
enable mass balance transfers within the pasture and livestock models.
Priority is given to higher quality pasture components when sheep
grazing is simulated, corresponding with their selective grazing habits.
As a result, most of the pasture (75%) was consumed from the 70 and
80% digestible pools. In the lower quality pastures 15–20% of intake
was from the 30–50% digestible pools. Total intake was higher for le­
gumes with higher DMD profiles (bladder clover and subterranean clo­
ver). Intake of pasture averaged over the year was 567 g/head/day
(Table 7).
The economic cost of the provision of supplementary feeding to
sheep varied by site and legume species grown (Fig. 7; checked bars).
For subterranean clover and bladder clover pastures, the cost of sup­
plementary feeding was consistently lower across all sites. At Cascade
and Condobolin there was a 28% lower cost of feeding with subterra­
nean clover and bladder clover pastures compared with the average of
other pasture legumes. There was little effect of legume species on
supplementary feeding costs among the remaining four legumes. Simi­
larly, predicted gross margins were AU$35/ha (or 15%) higher on
average in livestock enterprises where subterranean clover and bladder
clover were grown. This increase in gross margin was greater than the
saving in supplementary feeding costs, which was the result of higher
revenue from the livestock enterprise. This suggests other benefits of the
fertility or better rates of survival.
4. Discussion
The differences in the rate and level of decline in Dry Matter Di­
gestibility among six annual legume species was well represented in
calibrated Beta models using GrassGro™. In several instances, where
measured DMD was substantially higher than predicted, it was likely
related to the indeterminant growth patterns of some species extending
the growing period in these swards. Effects of pasture digestibility
modelled lamb growth rates were comparable with published research
(Fig. 8), albeit that lamb growth varied widely due to other factors such
as breed differences. This allowed a comparison of their productivity
and associated economic outcomes as pastures in a self-replacing Merino
sheep enterprise, at seven locations across southern Australia. Our hy­
pothesis that nutritive value was of greater economic importance than
higher biomass production in annual pasture legumes was supported by
the modelling results. Bladder clover and subterranean clover had the
highest nutritive value of the six annual legumes and provided the
highest feeding value to livestock, despite having lower net primary
productivity than some other species. The gross margin of the livestock
enterprise when bladder clover and subterranean clover were grown
increased by AU$35/ha compared with an average of the other species.
This is consistent with research in perennial species where economic
gains from genetic improvement of forage quality have been demon­
strated (Monjardino et al., 2014; Wims et al., 2017). This research
highlights the value of considering the nutritive value of annual pasture
species in their prioritisation for use in the temperate pasture systems of
southern Australia.
The conditions of our study assumed successful and persistent
establishment of the pastures, and that pastures were free of other bio­
logical constraints such as poor adaption to soil, sub-optimal nutrition,
variation in grazing management, plant sensitivity to heavy grazing.
Some of these constraints in GrassGro™ representing the potential
challenges in pasture establishment are described in more detail by
Clark et al. (2000). In practice, these factors would produce greater
variability in pasture production and result in biomass produced being a
more substantive economic driver, was the thesis of the study by

Fig. 4. Predicted net primary production (kg/ha) of six annual legume Beta models simulated under grazing across 30 years (1989–2018) for seven sites in the mixed
farming region of southern Australia. Error bars are standard errors of the means (n = 30).

7
D.T. Thomas et al. Agricultural Systems 194 (2021) 103272

80

70

60

50

Dry Maer
Digesbility 40
(%)

30

20

10

0
Barrel Medic Biserrula Bladder Clover French Strand Medic Sub Clover
Serradella
Fig. 5. Dry Matter Digestibility (%) of the season average green (filled) and immediately senesced (open) above-ground biomass for simulated Beta models for six
monoculture annual legume pastures, averaged across all seven sites. Error bars are standard deviations of the means (n = 30). (For interpretation of the references to
colour in this figure legend, the reader is referred to the web version of this article.)

Fig. 6. Proportion of total metabolisable energy intake in a simulated self-replacing sheep enterprise contributed from Pasture (filled), Stubble (dotted), Shrub
(checked) and Supplement (open) for six monoculture annual legume pastures modelled at seven sites in the mixed farming region of southern Australia.

8
D.T. Thomas et al. Agricultural Systems 194 (2021) 103272

Table 7 and seed-set are important for the adoption and return on investment
Predicted average intake (g DM/head/day) of pasture by ewes from different from new annual pasture legumes.
Dry Matter Digestibility (%) pools for six Beta pasture models simulated in The legume-based pastures simulated in our study were grown as
GrassGro™. monocultures for the purpose of allowing a clear comparison. However,
Pasture Dry Matter Intake of pasture of each digestibility Total in mixed swards the overall nutritive value of species may be more
pool (g/head/day) consequential but also interact with grazing intensity. That is, lower
30% 40% 50% 60% 70% 80% grazing intensity gives livestock greater opportunity to selectively graze
Barrel medic 19 25 32 42 159 247 524
quality plant species. Typical annual pastures of southern Australia tend
Biserrula 7 17 93 48 217 190 573 to be mixed swards of legumes, grasses and forbs, and the non-legume
Bladder clover 1 3 15 125 197 280 621 species tend to be relatively lower in nutritive value as dry feed (McI­
French serradella 15 60 27 37 209 197 545 vor and Smith, 1973; Doyle et al., 1989; Thomas et al., 2011). However,
Strand medic 12 32 37 45 163 251 540
annual pasture species may also form part of a mixed pasture integrated
Subterranean clover 3 12 82 62 193 248 599
Mean 9 25 48 60 190 236 567 with drought-tolerant perennial species to take advantage of sporadic
summer rainfall in these regions and used tactically during seasonal
periods of feed deficit has been demonstrated in other research (e.g. Bell
Sollenberger and Vanzant (2011). Sollenberger and Vanzant (2011) et al., 2008a; Norman et al., 2010; Nicol et al., 2013). Our results
found that pasture nutritive value sets the upper limit for livestock highlighted the substantive differences in nutritive value that exist
growth, the rate of decline in growth with increasing stocking rates and among annual legumes, and particularly in the rate of decline in nutri­
the level of pasture production at which the livestock growth begins to tive value across the season. For example, the DMD of strand medic
decline. Failure of pastures to establish is a major constraint to the decreased at an average rate of 0.30%units/day dry matter DMD from
adoption of new pasture legumes and so is an important consideration day 100 onward, compared with 0.10%units/day for bladder clover.
when choosing species for pasture improvement. This may diminish the Both the overall and seasonal profile of nutritive value in annual le­
importance of selecting species to improve sward nutritive value. gumes are key contributors to the differences in productivity of these
Compared with crops, annual pasture legumes are relatively small pastures.
seeded, less widely adapted to soil types, rely on regenerating from a Ultimately, the measure of success in annual pasture legumes is that
seed-bank, are typically grown with competing pasture species and often they contribute a high proportion of the annual feedbase. This measure
given lower management priority, all of which can reduce their capacity is consistent with previous research where the magnitude of the seasonal
to establish and persist in the face of variable climates and unpredictable feed deficit is considered, i.e. the marginal value of the provision of feed
seasons (Hackney et al., 2008; Hogg and Davis, 2009; Nichols et al., at different times during the season (Flugge et al., 2004). In the current
2012; Flohr et al., 2021; Nutt et al., 2021). Pastures species that fail to study, the proportion of Metabolisable Energy Intake from the annual
establish result in less productive pastures and impact the overall legume pastures relative to all other feedbase components varied by up
feedbase, with the additional cost of the initial investment to be recov­ to 10 percentage units within the same location. Although differences in
ered. In one study, the number of years to recover positive cumulative pasture quality have been shown to relate to differences in sheep
net cash flow was increased by between 14% to greater than 50% growth, these relationships may be quite variable given other factors,
(depending on stocking rate and pasture system) after a failed perennial particularly animal class and genetics (Fig. 8). These differences relate to
pasture establishment (Lewis et al., 2012). Addressing the constraints to the amount, quality and timing of pasture, or harvestable energy from
persistence by promoting best-practice management at establishment this forage, in relation to the seasonal energy demand of the livestock

Fig. 7. Gross margins (open, AU$/ha) and supplementary feeding costs (checked, AU$/ha) in a simulated self-replacing sheep enterprise for six annual legume
pastures modelled at seven sites in the mixed farming region of southern Australia.

9
D.T. Thomas et al. Agricultural Systems 194 (2021) 103272

Annual pasture legumes, produced through the domestication of

‘wild’ relatives - primarily of Mediterranean origin, require a wide range
of traits to be successfully integrated into the farming region of southern
Australia and a sophisticated understanding of existing farming systems
(Howieson et al., 2000; Ewing, 2017). In addition to improving the
feedbase, they must also fit in with cropping rotations, establish reliably
under difficult and variable conditions, and persist as regenerating
pastures unless they can be regularly re-established as a pasture phase in
a way that is economic (Revell et al. 2012). As a result, pasture breeding
objectives tend to be derived quite differently from the methods that
may be applied in animal breeding, where profit-based breeding indices
are common (Smith et al., 2014). Estimating the value of pasture
improvement is more straight-forward in permanent pastures, devel­
oped for areas where persistence is reliable even if susceptible to
degradation in composition (e.g. Wims et al., 2017). Selecting annual
legume genotypes that meet profit objectives for livestock businesses is
challenging and the importance of nutritive value may be overlooked,
particularly as these plants are typically preferred by livestock and of
higher in nutritive value than other forage plants. This study reiterates
the importance of systematically measuring nutritional value, in parallel
Fig. 8. Relationship between pasture quality (Digestibility %) and the growth to agronomic traits, in forage improvement programs. In future, inex­
rate of lambs in grazing of Masters et al., 2006 (x), McGrath et al., 2015 (●),
pensive and frequent measurement of nutritional value of a broad range
Norman et al., 2013 (■), Thompson et al., 2010 (Δ), Thomas et al., 2015 (+).
of forage species through NIRS analytical methods should help guide
Dashed line is GrassGro™ modelled data taken from the current study.
genetic improvement programs for new forage species (Norman et al.,
2020). Based on the results of our study, we make the case that the
(Bell et al., 2008b).
nutritive value of annual pasture legumes be considered, at least to the
The GrassGro™ model is a detailed and effective biophysical model
same extent as biomass production, in improving the role of pasture
for comparing grazing systems scenarios, which requires highly detailed
legumes in mixed farming businesses.
input processes for parameterising, calibrating and validating new plant
models. In this study we built Beta models to represent each of the
5. Conclusion
pasture species, which reflected differences in rates of growth and Dry
Matter Digestibility. Therefore, comparisons among the pastures was
There is an opportunity to increase the efficiency of livestock pro­
limited to this extent. For example, N content of the plant models was
duction on temperate pastures by exploiting the high variability in
not modified from base models, and genotypic differences in N content
nutritive value among annual pasture legume species. Historically this
that exist among the legumes would likely to affect their grazing value,
has been slowed by the many potentially competing objectives in
particularly in senescent material (Thomas et al., 2010a). Where sheep
pasture improvement, and any new candidate species will still need to
were grazing pasture legume monocultures this is unlikely to be an issue,
accommodate the broader role of pasture legumes in farming systems.
since both modelled and observed N content are typically higher than
This study highlights the need for plant nutritive value across all
sheep requirements at this time. However, in mixed pastures containing
phenological stages to be at the forefront of traits considered when
species with relatively low N content there would have greater conse­
selecting and developing annual pasture legume species. Failure to do so
quence for livestock production, particularly for young animals. In this
will likely result the production of biomass that is sub-optimal to support
study, access to data to calibrate the growth of the indeterminant
animal production.
pasture species, such as biserrula or serradella, in different seasons
Supplementary data to this article can be found online at https://doi.
under different levels of water stress was limited. Therefore, effects of
org/10.1016/j.agsy.2021.103272.
differences in the amount and quality of pasture available in the
potentially much longer growing periods of indeterminate pasture le­
Declaration of Competing Interest
gumes were not fully investigated. However, this growth habit was able
to be represented within the model by integrating perennial character­
None.
istics from the white clover model. Semi-determinant growth potentially
extends the growing season of some annual legumes, where soil mois­
Acknowledgements
ture is available, but provides the added advantage of producing seed
over an extended period, which helps ensure to maintain a seed-bank
The authors acknowledge The Dryland Legumes Pasture Systems
and pasture regeneration (Loi et al., 2014). However, the value of this
project, which is funded by the Australian Government Department of
difference in growth habit needs to be demonstrated in a systems
Agriculture Water and the Environment as part of its Rural R&D for
context, in relation to the timing of the available feed. Apparent benefits
Profit program, the Grains Research and Development Corporation,
of a feed component can be much higher than the actual value of the
Meat and Livestock Australia and Australian Wool Innovation (Project
component as part of the overall feedbase (Thomas et al., 2010b). This is
No. RnD4Profit-16-03-010). The research partners include the South
due to the timing of when feedbase components are available, and dif­
Australian Research and Development Institute, Murdoch University,
ferences in the marginal value of feed at different times of the year
the Commonwealth Scientific and Industrial Research Organisation, the
(Flugge et al., 2004; Bell et al., 2008b). Based on these considerations,
WA Department of Primary Industries and Regional Development, NSW
and the season to season variability in pasture growth, biophysical
Department of Primary Industries and Charles Sturt University, as well
modelling of pasture systems arguably provides the best option to ensure
as grower groups. We gratefully acknowledge model calibration data
new plant and animal models can be applied as accurately and effi­
from ‘Efficient Livestock and Low Emissions (ELLE) from southern
ciently as possible. Overall, the GrassGro™ model provided an effective
grazing systems’ funded through the Australian Department of Agri­
method to compare effects of changes in plant nutritive value within a
culture and Water Resources ‘Filling the Research Gap’ programme and
grazing enterprise.
managed as part of the National Livestock Methane Programme by Meat

10
D.T. Thomas et al.
and Livestock Australia (project Number: 01200.042; B.CCH.6540), a
collaboration between the University of Western Australia (UWA),
South Australian Research and Development Institute (SARDI) and the
Commonwealth Scientific and Industrial Research Organisation
(CSIRO).
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