doi:10.1017/S1742170510000347
Renewable Agriculture and Food Systems: 26(1); 72–91
The potential of herbaceous native
Australian legumes as grain crops: a review
Lindsay W. Bell1*, Richard G. Bennett2, Megan H. Ryan2, and Heather Clarke3,y
1
CSIRO Sustainable Ecosystems, PO Box 102, Toowoomba, Qld 4350, Australia.
School of Plant Biology M084, University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia.
3
Centre for Legumes in Mediterranean Agriculture (CLIMA), University of Western Australia, 35 Stirling Highway,
Crawley, WA 6009, Australia.
*Corresponding author: Lindsay.Bell@csiro.au
2
Accepted 25 June 2010; First published online 18 August 2010
Research Paper
Abstract
Many agricultural systems around the world are challenged by declining soil resources, a dry climate and increases in
input costs. The cultivation of plants that are better adapted than current crop species to nutrient poor soils, a dry climate
and low-input agricultural systems would aid the continued profitability and environmental sustainability of agricultural
systems. This paper examines herbaceous native Australian legumes for their capacity to be developed as grain crops
adapted to dry environments. The 14 genera that contain herbaceous species are Canavalia, Crotalaria, Cullen, Desmodium,
Glycine, Glycyrrhiza, Hardenbergia, Indigofera, Kennedia, Lotus, Rhynchosia, Swainsona, Trigonella and Vigna. A number
of these genera (e.g., Glycine, Crotalaria, Trigonella and Vigna) include already cultivated exotic grain legumes. Species
were evaluated based on the extent to which their natural distribution corresponded to arid and semi-arid climatic regions, as
well as the existing information on traits related to harvestability (uniformity of ripening, propensity to retain pod, pod
shattering and growth habit), grain qualities (seed size, chemistry, color and the absence of toxins) and fecundity. Published
data on seed yield were rare, and for many other traits information was limited. The Australian species of Vigna, Canavalia
and Desmodium mainly have tropical distributions and were considered poorly suited for semi-arid temperate cropping
systems. Of the remaining genera Glycyrrhiza and Crotalaria species showed many suitable traits, including an erect growth
habit, a low propensity to shatter, flowers and fruits borne at the end of branches and moderate to large seeds (5 and 38 mg,
respectively). The species for which sufficient information was available that were considered highest priority for further
investigation were Glycine canescens, Cullen tenax, Swainsona canescens, Swainsona colutoides, Trigonella suavissima,
Kennedia prorepens, Glycyrrhiza acanthocarpa, Crotalaria cunninghamii and Rhynchosia minima.
Key words: domestication, novel crops, perennial, adaptation, arid, climate change
Introduction
Increasing crop diversity can reduce our reliance on just a
few major food crops and improve the sustainability and
resilience of agriculture in the future1,2. With dry climatic
conditions, reduced allocations of water for agriculture and
increasing demands for food production from currently marginal areas, species adapted to more stressful environments
are needed. In addition, alternative crops with improved efficiency of fertilizer use and reduced reliance on pesticides
would improve the sustainability of our agricultural
yPresent address: The University of Notre Dame, PO Box 1225,
Fremantle, WA 6959, Australia.
systems3. Benefits of protection of soil from erosion, reduced leaching of water and nutrients, and additional forage
for livestock could also be provided by perennial grain
crops2,4,5. Exploring the wild native flora provides an exciting and substantial opportunity to identify species with
potential as alternative grain crops for the future6.
Australia, because of its arid climate and infertile and
poor soils, is a good place to look for potential new grain
crops adapted to harsh growing environments. Yet, the
potential of Australia’s native flora for use in agriculture
has been relatively underexplored. Some Australian grasses
and legumes have been investigated as potential pastures
or forage species7–13, but little work has been conducted
on their suitability as grain crops. Woody legumes such as
Acacia spp. could have some use as alternative sources of
# Cambridge University Press 2010
Potential of native Australian legumes as grain crops
grain14, but herbaceous species are more suited to modern
broad-acre farming systems because they can be mechanically harvested and are more easily removed and rotated
with other crops. One Australian grass, Microlaena
stipoides, has been investigated to a limited extent for
grain production15, yet herbaceous legumes have received
little attention. Rivett et al.16 examined a number of native
Australian plants for their potential as grain crops and
found that the legumes Hardenbergia violacea, Crotalaria
cunninghamii and Kennedia nigricans warranted further
examination as they possessed relatively large seeds with
substantial amounts of crude protein and oil.
There are a few modern examples where efforts have
been made to domesticate legumes for grain production in
agricultural systems where grain legumes are/were lacking;
Lupinus angustifolius (narrow-leafed lupin)17, Lupinus
luteus (yellow lupin)18 and Desmanthus illinoensis (Illinois
bundleflower)19. Evidence with these species and advances
in our understanding of crop domestication and in the technologies associated with crop breeding should allow rapid
advances in the future20. However, the domestication of
Australian legumes may be more difficult, as there is little
or no history of predomestication. This means that the net
may need to be cast wide, as many species are unlikely
to possess traits common to domesticated plants21. While
Australian aboriginals manipulated their environment to
ensure food supply, notably through the use of fire, they did
not generally practice agriculture in a way close to modern
cultivated cropping systems. In addition, while seed
grindstones have been found in many areas and there are
reports of aboriginal seed collecting from grasses and of the
use of seed from 50 species of Acacia, there is no
indication that the seeds of native herbaceous legumes were
other than a very occasional source of food22. Hence,
Australian native herbaceous legumes have not been subjected to the same predomestication pressures that have
acted upon other species that have been cultivated by ancient peoples or were simply present (as weeds) in early
agricultural systems23,24.
We examined the 14 genera of Australia’s native
legumes that contain herbaceous species for their suitability
as grain crops. While little useful information was available
for many species, we found that at least nine species merit
further investigation. Species identified were most likely to
be adapted to the climate of Australia’s semi-arid cropping
regions, but they may have applications in other semi-arid
environments throughout the world or in areas predicted to
experience a dry climate in the future.
Approach and Desirable Plant Attributes
Among Australia’s legumes, there are 14 genera that contain herbaceous species. Information was gathered on three
main aspects: species distribution in arid or semi-arid climatic range; traits related to harvestability, grain size and
yield potential; and grain chemistry and nutritional qualities
(discussed below). Some genera also include currently
73
cultivated grain legume crops exotic to Australia (e.g.,
Glycine, Vigna, Trigonella and Canavalia). This close relationship could indicate genera that possess suitable agronomic characteristics, or closely related species that may be
suitable for hybridization with the cultivated crop, to either
improve the agronomic traits of a wild species or transfer
desirable characteristics into the cultivated species (e.g.,
abiotic or biotic stress tolerance)5. The hybridization of
Australian perennial Glycine species with soybean is one
such example25,26. Together, this information is used to
identify genera and species with the most desirable attributes and the greatest immediate potential as grain crops.
Information was not available for some aspects of some
species, particularly in rarer or less studied species. Hence,
suppositions were drawn only where sufficient information
was available. Other species may also have desirable
characteristics or potential in different agro-climatic conditions. Beyond the scope of this review was an assessment
of the weed risk of these species. Indigenous species can be
regarded as weeds when growing outside their natural
range, and some species of Fabaceae are commonly mentioned in this context in Australia including some of the
genera assessed in this paper27.
Potential adaptation to arid and
semi-arid environments
Information on the distribution of Australian native herbaceous legumes was obtained from collection locations
available from the Australian Virtual Herbarium28 and
matched against Australia’s agro-climatic regions29. Species were prioritized if their distribution corresponded to
the arid interior (G and E6) or semi-arid environments with
sufficient capacity for plant growth in winter–spring (E2,
E3 and E4) (Fig. 1). Species that occur in these agroclimatic regions likely possess adaptations to short or
erratic growing seasons, and hot and dry climatic conditions, such as physiological drought tolerance mechanisms or reproductive strategies which enable them to avoid
these stresses (e.g., rapid flowering and deep roots).
Excluded from the target region were tropical (i.e., H, I1,
I2, I3, J1, J2 and E7) and cold climatic (i.e. B1 and B2)
regions, because plant growth is limited during winter–
spring due to lack of moisture and cold, respectively. Also,
Australian species from northern origins are more likely to
have flowering promoted by short days (short-day plants),
while long-day plants grow at higher latitudes, including
the Australian agricultural zone. Short-day plants have limited potential in these southern temperate areas. Although
cool-season grain crop production is common in agroclimatic regions E1 (wet ‘Mediterranean’) and D5 (coolseason and wet), the target region was restricted to the
less favorable climatic regions with a shorter winter–spring
growing season (i.e., E2—dry ‘Mediterranean’, E3—
temperate, subhumid and E4—subtropical, subhumid).
Agro-climates F3 and F4 are warm and wet environments
and were also excluded as they have a few climatic stresses
74
L.W. Bell et al.
Figure 1. Agro-climatic regions of Australia (adapted from29). The target climatic regions include the arid and semi-arid regions too dry
for field crops (i.e., G and E6) and the semi-arid cropping zone where moisture is a major growth limitation and with sufficient capacity
for growth in winter–spring (i.e., E4, E3 and E2).
that reduce plant growth throughout the year. A wider
species distribution was also regarded as favorable as it
suggests greater adaptability and a greater capacity to exploit within species variability.
Harvestability
Plant traits that influence grain harvestability are critical
in the domestication process30 and hence were considered
important aspects for evaluating the agronomic potential
of wild legume species. Plants with a self-supporting, erect
or semi-erect growth habit and those that set pods close
to the top of the plant would be most favorable for mechanical harvesting, while highly prostrate species may be
difficult to harvest. Species with a twining or rambling
habit were not regarded as ideal, but were not removed
from consideration. Many current grain legume crops
originated from ancestors with a climbing, creeping or
straggling growth habit and their domestication has shortened internode length and reduced indeterminate branching
(e.g., Phaseolus, Vigna, Glycine, Pisum and Arachis)31.
Pod dehiscence and indeterminate growth habit is ubiquitous among undomesticated legumes and domesticated
grain and forage legumes alike (e.g., soybean32 and birdsfoot trefoil33). While pod indehiscence (non-shattering)
overcomes some agronomic challenges, it is clear that
this is not a disqualifying trait, as selection for indehiscence
during domestication has occurred for most modern crops
and would also occur in native Australian legumes.
Grain size and yield potential
Legume grain or seed size is obviously an important aspect
as it influences the potential market uses and agronomic
performance. Large seeds also offer advantages for crop
establishment especially from greater depth, under greater
competition (e.g., weed burden) and in low-nutrient or
moisture conditions34. Cultivated grain legumes have large
seeds compared to their wild relatives and seed size has
been increased substantially through active selection, and
hence it is likely that the seed size of wild legumes would
be smaller than that of cultivated grain legumes. For
example, the seed size has increased at least tenfold in
Phaseolus coccineus (French bean) and by at least fivefold
in other legume species31. This is also demonstrated in the
germplasm of L. angustifolius (narrow-leaf lupin), where
seed size may vary substantially from 29 to 244 mg, with
the ‘wild’ types generally smaller seeded35. Despite the
appeal of species with larger seeds, small seeds may actually be equally appealing, especially if they contain high
concentrations of a desirable product such as oils (e.g.,
Brassica napus, canola). Attractive small seeds or those
that have special properties or novel appearance may also
have a market as whole grains similar to sesame (Sesamum
indicum), poppy (Papaver somniferum) or linseed (Linum
usitatissimum).
Species exhibiting high overall fecundity and the capacity to self-fertilize are highly desirable. Most domesticated
grain legumes are self-fertilizing with the exception of
P. coccineus, Vicia faba and Cajanus cajan31. Some difficulties might occur with domesticating outcrossing species
Potential of native Australian legumes as grain crops
due to the ability to outcross with their wild counterparts
and to reintroduce undesirable traits. Species capable of
self-fertilizing would be less problematic for future breeding. Annual species may have a greater overall fecundity,
because their survival relies on producing viable seeds,
but perennial species may be equally productive provided
they flower and reproduce in their first year5. Many domesticated annual grain legumes have originated from a
perennial life form, most likely because of the selection
pressure for increasing seed yield31.
Grain chemistry and nutritional qualities
Native legumes found to produce seeds with high concentrations of protein and/or oils/fats are clearly desirable. In
addition, those with favorable amino acid or fatty acid profiles or the presence of unique compounds that can benefit
human health may have a significant market as a health
food. In most cases, little information is available on the
nutritive qualities of native Australian legumes. On the
other hand, a number of Australia’s native legumes are
known to possess potent bioactive compounds, some of
which may be toxic (e.g., swainsonine and hydrogen
cyanide (HCN)), but some of which have pharmaceutical
functions or can provide human health benefits at the
correct concentrations (e.g., furanocoumarins and phytoestrogens)36,37. Many cultivated grain legumes contain antinutritional compounds which have been lowered by
breeding (e.g., alkaloids in lupins)38.
Canavalia
The genus Canavalia consists of approximately 70 species
mostly of tropical origin. Several species are legume grain
crops of secondary importance, including common jackbean (Canavalia ensiformis), sword bean (Canavila gladiata) and Canavalia cathartica. Raw seeds of Canavalia
contain a number of anti-nutritional factors including
phenolics, tannins, saponins, concanavalin A, canavanine,
cyanogenic glycosides and HCN39,40. Canavalia are famous for the presence of the lectin, concanavalin A which
has commercial importance as a reagent in glycoprotein
biochemistry and immunology41,42. Four species of Canavalia are found in Australia but none are endemic;
Canavalia rosea, Canavalia carthartica, Canavalia sericea
and Canavalia papuana. These are mostly found in tropical, coastal hinterland regions (Fig. 2a). While C. rosea is
found further south than other species into the subtropics, it
is mainly confined to coastal and high-rainfall areas
(Fig. 2a). Because Canavalia match poorly with our target
climatic regions, they are not considered further here,
although they may have some potential as an adapted
legume crop for the tropics. They possess large seeds and
are a rich protein source39,42.
75
Crotalaria
Crotalaria is a genus of herbaceous plants and woody
shrubs commonly known as rattlepods because seeds become loose in the pod as they mature and rattle when the
pod is shaken. Around 600 or more species of Crotalaria
are described worldwide, mostly from the tropics with at
least 500 species known from Africa; 19 species are native
to Australia43. Some exotic species of Crotalaria have
agronomic uses (e.g., Crotalaria spectabilis, Crotalaria
ochroleuca, Crotalaria longirostrata and Crotolaria juncea
(sunn hemp))43.
The Australian native Crotalaria species are mainly
found in tropical regions. Four species occur further south
in the target region; Crotalaria eremaea (desert rattlepod),
Crotalaria mitchelli (yellow rattlepod), C. cunninghamii
(green birdflower or parrot pea) and Crotalaria dissitiflora
(plains rattlepod) (Fig. 2b). C. eremaea and C. cunninghamii occur mainly on sandy or well-drained soils in lowrainfall regions of central Australia44. C. mitchelli occurs
on sandy soils in the tropical and subtropical areas of
the east coast with > 500 mm mean annual rainfall45.
C. dissitiflora occurs on heavy clay soils also in the subtropics and tropics although it occurs further inland and in
lower-rainfall regions than C. mitchelli45.
Crotalaria includes annual, biennial and perennial species that can range in form from herbs to shrubs (0.3–3 m
high). The four species occurring in the target region have
erect or ascending habits; C. cunninghamii is an erect
perennial subshrub growing to 1 m or higher, C. eremaea
is an erect subshrub 0.5–1 m high, C. dissitiflora is an
erect-sprawling short-lived perennial <30 cm high and
C. mitchelli is an erect—decumbent woody forb about
60 cm high44. All these species flower in winter–spring
(C. cunninghamii sometimes in autumn) and are generally
open pollinated by insects. C. dissitiflora has been observed
to shed its leaves during winter45. A notable and advantageous characteristic of these species is that flowers and
pods are borne at branch ends46, which favors mechanical
harvesting. Some Crotalaria shatter explosively, while
pods of others are more stable and slower to shatter46.
Some Crotalaria have large seeds (e.g., 38 mg in
C. cunninghamii) (Table 1), while others are smaller
(e.g., those of C. dissitiflora are only 2–3 mm long). Seeds
are often smooth and vary in color (yellow in C. dissitiflora,
greenish–gray in C. mitchelli and red–brown in Crotalaria
smithiana). Due to its large seed size and substantial protein
and oil content (Table 2), C. cunninghamii was previously
identified as a species worthy of further investigation16.
Toxic pyrrolizidine alkaloids are produced by some members of this genus and these can be poisonous to livestock47,
but whether these are present in seeds of the four species
that occur in the target region is unknown. C. dissitiflora is
suspected of poisoning livestock, but there is conflicting
evidence45. Everist47 suggests that the toxicity might be
lost when plants are cut. C. eremaea is often eaten by
sheep, suggesting low or no alkaloid problems and
76
L.W. Bell et al.
(a)
(c)
(b)
(d)
Figure 2. Distribution of a selection of widely distributed Australian native Canavalia (a), four native Crotalaria spp. (b) and Cullen spp.
(c, d) mapped against targeted agro-climatic regions; semi-arid cropping zone (E2, E3 and E4)—light gray; and the arid interior (E6 and
G)—dark gray (see Fig. 1). Data sourced from Australian Virtual Herbarium28.
C. cunninghamii is reputedly edible by humans without any
indication that prior treatment is necessary44,48.
Overall, Australian Crotalaria seem to have a number of
characteristics which suggest they warrant further investigation for their potential as grain crops. In particular,
C. cunninghamii has a desirable growth habit, produces
large seeds which contain high levels of protein and
some oil, and does not seem to produce toxic alkaloids
(Table 3). Little agronomic information was available
on the other three species found in arid and semi-arid
regions of Australia (i.e., C. eremaea, C. mitchelli and
C. dissitiflora), but these may also warrant further
investigation.
Cullen
The Cullen genus includes 32 species of which 25 are endemic to Australia49. It has been explored for forage plants
in the past and again recently in Australia11,50,51. While no
species of Cullen are used commercially, the closely related
Psoralea genus includes one economically important plant
native to India. Psoralea corylifolia seeds have medicinal
properties, which are thought to be imparted due to their
content of furanocoumarin, in particular psoralen.
Cullen in Australia is widely distributed across a range of
climates from summer- to winter-dominant rainfall and the
average annual rainfall of the distribution across species
ranges from 200 to 1300 mm51. All species of Cullen occur
within the target region and 12 of these species mainly
occur in low-rainfall environments with an annual average rainfall £ 400 mm51. Of these species, Cullen australasicum, Cullen graveolens, Cullen pallidum and Cullen
discolor mainly occurred in the lower-rainfall regions
(Fig. 2c and 2d). Cullen cinereum has a slightly more tropical distribution than the other species; although, it is also
found throughout the target zone (Fig 2c). Cullen species
have been reported to have excellent drought tolerance
where they have been evaluated as forage plants11,52.
All species have a deep tap-root which may become woody
in the perennial species. Roots of Cullen patens
Potential of native Australian legumes as grain crops
77
Table 1. Seed size of Australian native herbaceous legumes.
Means in brackets, where a range of material has been measured.
Species
Seed mass (mg)
Crotalaria cunninghamii
Cullen australasicum1
C. cinereum1
C. cinereum1
C. cinereum1
C. graveolens1
C. pallidum1
C. patens1
C. patens1
C. plumosum1
C. tenax1
Glycine canescens
G. canescens
G. clandestina
G. tomentella
G. latifolia
G. latifolia
Glycyrrhiza acanthocarpa
Hardinbergia violacea
H. violacea
H. comptoniana
Indigofera colutea
I. linnaei
I. linnaei
I. australis
Kennedia coccinea
K. coccinea
K. eximia
K. nigricans
K. prorepens
K. prorepens
K. prostrata
K. prostrata
K. prostrata
K. rubicunda
Lotus australis
L. australis
L. cruentis
Rhynchosia minima
R. minima
R. minima
R. minima
Swainsona canescens
S. canescens
S. colutoides
S. kingii
S. formosa
S. purpurea
S. swainsonoides
Trigonella suavissima
Vigna radiata ssp.
sublobata
V. lanceolata
V. vexillata
V. vexillata
38.5
5.5
4.2
4.1
5.5
2.3
3.2–6.4 (4.8)
8.4
2.82
11.3
4.4
5.6
5.9–8.9
4.2
5.0
11.1
6.6–12.5
5.4
38.5
22
38.3–45.2
1.1
1.8
1.9
5.2
8.8
26.3
9.0
15.6
12.4
6.6
29.7
31.9
9.3–44.4
24.4
2.7
1.3–1.9
1.8
11.8
8.8–20.4 (12.1)
9.4
16.8
2.1
1.3
2.9
2.6
3.9
2.8
7.2
1.0
7.4–27.0 (13.4)
1
2
20–34
8.1–17.7
7.0–19.5 (11.1)
Indicates whole fruit (pod + seed).
Immature seeds harvested.
Reference
16
11
108
36
Unpublished
Unpublished
Unpublished
108
57
36
Unpublished
108
Unpublished
109
57
110
69
Unpublished
16
109
111
108
108
Unpublished
16
16
111
Unpublished
16
112
Unpublished
111
86
Unpublished
109
112
Unpublished
88
57
91
108
Unpublished
108
Unpublished
Unpublished
Unpublished
Unpublished
Unpublished
Unpublished
Unpublished
104
103
113
105
data
data
data
data
data
data
data
data
data
data
data
data
data
data
data
data
data
data
data
(syn. Psoralea eriantha) have been reported to penetrate to
a depth of 4 m, and this was associated with the drought
resistance of this species53. Another evident adaptation
to drought is the dense coverings of glandular hairs on
the leaves of some Cullen species (e.g., C. pallidum and
C. patens). Strong soil-type associations are rare in many
Cullen species (e.g., C. australasicum), but some species
have particular preferences; for example, C. tenax seems to
prefer heavy clay soils and C. pallidum is predominately
found on deep sand dunes and sandy soils45,49,51.
The Australian species of Cullen include shrubs, subshrubs and herbs and a number of them have a favorable
growth habit and phenology, as outlined below. Nineteen
species are herbaceous or semi-herbaceous, of which 16 are
perennial or short-lived perennials. C. graveolens, Cullen
plumosum and Cullen walkingtonii are annual or biennials49. Most Australian taxa bloom in the first year49.
Flowering mainly occurs in spring, but indeterminate
flowering continues throughout the year provided sufficient
moisture is available49,53. In a glasshouse study, Bourgaud
et al.36 recorded that flowering occurs around 40 days after
germination in C. cinereum (about 900 degree days) and
around 60 days after germination in C. plumosum (1340
degree days). Flowering of C. australasicum and C. patens
is controlled by day length according to Britten and
De Lacy54, with long-day treatments (i.e., < 12–13 h dark)
inducing flowering. They also found that genotypes vary in
their response, indicating differences in phenological
adaptations within these species. In C. australasicum,
flowering and fruiting times are extremely variable in the
first year, but with greater synchrony in the second year11.
Many Cullen also seem to be capable of self-pollinating.
Britten and Dundas55 found that erect types in the Psoralea
patens complex (i.e., C. australasicum) were 50–75% selfing, while the prostrate and semi-erect lines (i.e., C. patens)
were outcrossing only. Bourgaud et al.36 noted that
C. cinereum and C. plumosum are capable of selfpollinating. Using microsatellite markers, Kroiss et al.56
estimated the outcrossing rate in C. australasicum to be
at least 3–13% and hybrids were formed with C. pallidum,
but not C. discolor or C. patens.
A couple of studies have found Cullen to allocate
significant resources to reproduction and produce useful
amounts of seed. Bourgaud et al.36 found seed yields up to
1.65 g plant - 1 (47% of dry matter) from C. cinereum and
1.75 g plant - 1 for C. plumosum (60% of dry matter). The
higher yield from C. plumosum was due to the greater seed
mass (11.3 mg), while the whole plant biomass was less
than C. cinereum. The production of seeds from C. tenax
has also been measured at 22 g plant - 1 (4820 seeds)
(Bennett, unpublished data). Kerridge and Skerman53 recorded that the reproductive parts of C. pallidum made up
42% of plant biomass when the plants were left to grow for
12 weeks.
Cullen are characterized by indehiscent (non-shattering)
fruits with the seeds adherent to the pericarp (pod). Fruit
sizes of Cullen typically range from 4 to 6 mg, although
78
L.W. Bell et al.
Table 2. Chemical composition of seeds of some native Australian legumes16. Seeds were obtained from a commercial native seed
service and had presumably been collected from the wild.
Fatty acid composition (% of total fat)
Species
Crude protein1
(%)
Fat
(%)
16 : 0
18 : 0
18 : 1
18 : 2
18 : 3
P : S ratio
23.3
21.0
18.8
27.4
23.9
3.8
8.1
2.8
3.0
9.1
17.2
12.1
17.6
12.3
14.9
6.8
5.1
3.8
7.0
6.5
26.0
23.2
26.3
29.2
30.2
46.5
55.9
45.6
43.9
41.0
3.3
5.0
6.2
6.4
4.4
2.1
3.4
2.4
2.6
2.1
Crotalaria cunninghamii
Hardenbergia violacea
Indigofera australis
Kennedia coccinea
Kennedia nigricans
1
2
2
Protein was calculated as 5.7r%N.
Ratio of polyunsaturated to saturated fatty acids.
Table 3. Prioritization of species for further investigation as grain legume crops. Species with little information are not included
(?—indicates where information is unknown).
Species
Highest priority species
Cullen tenax
Crotalaria
cunninghamii
Glycine canescens
Glycyrrhiza
acanthocarpa
Distribution1
Life
cycle2
Habit3
for further investigation
****
P
Sp, C
****
P
E
Seed size4
Flowers in
first year
Pollination
Pod/seed
retention
Small
Large
Y
?
Selfing
?
Low
?
****
****
P
P
T
E, C
Mod.
Mod.
Y
?
Selfing
?
Low
High
Kennedia prorepens
Rhynchosia minima
Swainsona canescens
Swainsona colutoides
****
***
****
***
P
P
A/B
A/B
T
T–SE
SE, Sp
E
Mod.–Large
Mod.–Large
Small
Small
Y
?
Y
Y
Open
Selfing
Open
Selfing
Low
Variable
Mod-High
Mod, delayed
Trigonella suavissima
****
A
E–SE
Small
Y
?
?
Y
Y
Y
Y
Variable
Variable
?
Variable
Low
?
Moderate priority species with a number of
Cullen australasicum
****
P
Cullen cinereum
***
A/B
Cullen graveolens
***
A/B
Cullen pallidum
***
P
suitable attributes
E
Mod.
E, Sp
Small–mod.
E–SE
Small
Sp
Small–mod.
Glycine latifolia
**
P
T
Mod.
Y
Open
Selfing
?
Open, some
selfing
?
Glycine tabacina
Kennedia coccinea
Indigofera australis
Swainsona formosa
Swainsona
swainsonoides
**
*
***
****
***
P
P
P
A
P
T
P, T
E, Sp
SE, Sp
Sp
Small–mod.
Large
Mod.
Small
Small
Y
Y
?
Y
Y
?
?
?
?
Open
?
Low
N
?
Y
Open
Open
Selfing
Very low
Species with some valuable attributes but some limitations
Kennedia prostrata
**
P
P, T
Large
Kennedia nigricans
*
P
P, T
Large
Lotus cruentus
****
A
Sp, C
Small
1
2
3
4
Other notable
qualities/
information
Aerial seed
Pods on
branch ends
Salt tolerance
and pods easy
to thresh
Good pod
retention
Waterlogging
& potential
salt tolerance
Collected and
selected as
a forage
Match between species distribution and targeted agro-climates; ****—highly favorable, ***—favorable, **—moderate, *—poor.
P, perennial; A, annual; B, biennial.
E, erect; SE, semi-erect; P, prostrate; T, twining/trailing; Sp, spreading; C, clumping/crown forming.
Large, 10–20 mg; Mod., 5–10 mg; Small, < 5 mg.
Potential of native Australian legumes as grain crops
fruits >8 mg have been measured in C. patens and
C. plumosum (Table 1). Smaller fruits (2.8 mg) were found
for C. patens by Silcock and Smith57, but this included many
immature seeds which probably reduced the average seed
mass. The non-shattering nature of Cullen is advantageous
for harvesting, but fruit retention on the plant is variable.
Skerman58 reports that ripe pods of C. patens drop to the
ground and seed harvesting would need to be performed by
suction. The pods of several species (e.g., C. australasicum,
C. patens, C. pallidum and C. discolor) fall from the plant
enclosed in the calyx, which can be very hairy49. This, and
the adherence of the seed to its pod, also poses some
complications about the ability to thresh the seed of Cullen,
unless processing could utilize the whole fruit. Dear et al.11
state that the seed of C. australasicum is easily threshed
from the pod without damage. However, since the seed is
completely adhered to the pod, it is likely that they were
referring to the removal of the woolly calyx material.
No information was found on the protein or oil content of
Cullen seed/fruit, but like other members of the Psoraleae
family, Australian Cullen species are known to contain the
furanocoumarins psoralen and angelicin59–61. Furanocoumarins are potent photosensitizing agents that may cause
phototoxic reactions, but they are also pharmaceutically
useful for the treatment of skin disorders such as psoriasis,
vitiligo, leukoderma and leprosy62. The seeds of Australian
Cullen species have been found to contain between 1000
and 8000 mg kg - 1 dry weight (DW) of furanocoumarins
(depending on species) and have been proposed as potential
sources for pharmaceutical use36,61,62. C. cinereum (syn.
Psoralea cinerea) and C. plumosum were identified with
the highest levels of furanocoumarins, but they have also
been measured in Cullen lachnostachys (syn. Psoralea
lachnostachys) and Cullen pustulatum (syn. Psoralea
pustulata) and are likely to exist in many other species.
The fruits generally contain the highest concentration of
furanocoumarins, up to 5500 ppm in Cullen corylifolia
(native to India) and the majority ( >70%) of this is found
in the cotyledon of the fruit60,63. Vegetative material may
also contain significant levels of furanocoumarins (up to
1600 ppm)60,64, which may affect the health of grazing
animals by inducing photodermatitis. However, furanocoumarins also play an important role in plant health by
controlling pathogens and insect activity.
In addition to furanocoumarins, some Psoralea and Cullen
species can also contain the flavonoid, daidzein, which is
increasingly studied because of its activity in cancer
prevention and treatment65. Daidzein has been found in the
fruits of two Australian species, C. cinereum (8.2 mg g - 1
DW) and C. tenax (27.5 mg g - 1 DW) and was also present
in their stems61,65. C. patens and C. cinereum were found
to also contain lectins and trypsin inhibiting proteins39.
A number of Cullen species warrant further investigation
for potential as grain crops (Table 3), because they are able
to produce large amounts of seed of moderate size, are selfcompatible, have an erect growth habit, and the ability to
grow and flower in their first year. High-priority species
79
have a reputation for high seed production (e.g., Cullen
tenax and C. cinereum) or display an annual life cycle (e.g.,
C. graveolens). Also an exciting attribute is the likely presence of furanocoumarins which may provide a pharmaceutical market for Cullen seed. However, the problem of
separating the seed from the calyx is a significant issue and
their market success would rely on uses that can utilize the
whole fruit.
Desmodium
Desmodium, also known as tick-trefoils or tick clovers, is a
large and taxonomically confusing genus containing about
300 species of which 21 species are native to Australia43.
No Desmodium species are grown as grain crops, but some
are cultivated as forage for livestock (e.g., Desmodium
intortum and Desmodium uncinatum) and as living mulch
or green manures.
Most Australian Desmodium occur in the tropics and
subtropics and only three species are distributed within
the target region; Desmodium varians (slender ticktrefoil), Desmodium campylocaulon (creeping tick-trefoil)
and Desmodium brachypodum (large tick-trefoil) (Fig. 3a).
D. varians occurs at the furthest south in the temperate
regions of Australia, but is mainly found in moister regions.
D. varians is a trailing or twining perennial that can flower
all year round, although flowering is usually concentrated
in the warmer months. D. campylocaulon and D. brachypodum are mainly found in the inland subtropics and
tropics. Both are erect and twining long-lived perennial
subshrubs growing up to 60–100 cm high, which flower
from late spring to autumn46. All three species have a
warm-season-dominant growth pattern and seem to prefer
climates where rainfall is summer-dominant.
An advantageous characteristic of Desmodium species
is that their reproductive racemes are at branch ends and
in some species are held well above the foliage (e.g.,
D. brachypodum)46. Desmodium pods also do not split or
dehisce at maturity, but they have a saw-like pod with segments that separate at maturity which enclose an individual
seed (known as a loment) and are individually dispersed46.
No information was found on the seed constituents or the
presence of toxins or other bioactive compounds in the seed
of Australian Desmodium, although some Desmodium
species are known to contain alkaloids in their leaves.
Overall, we consider the Desmodium species to be of
marginal interest because of the little information on the
agronomic and seed attributes, and their tendency toward
moister- and summer-dominant rainfall environments.
Glycine
Australia is the center of diversity for the Glycine genus,
which contains the most important legume grain crop
worldwide, soybean (Glycine max). The 23 Glycine species
native to Australia make up the subgenus Glycine, while the
cultivated G. max and its ancestor Glycine soja make up the
80
L.W. Bell et al.
(a)
(b)
(c)
Figure 3. Distribution of Australian native Desmodium spp. (a)
and Glycine spp. (b, c) mapped against targeted agro-climatic
regions; semi-arid cropping zone (E2, E3 and E4)—light gray; and
the arid interior (E6 and G)—dark gray (see Fig. 1). Data sourced
from Australian Virtual Herbarium28.
subgenus Soja, which originates in South-East Asia.
Numerous attempts to hybridize wild Glycine and G. max
have been made and have mainly been successful with
tetraploid (2n = 80) types of Glycine tomentella5. Australian
Glycine species have been investigated for beneficial
traits for soybean improvement, such as drought tolerance66, 2–4D resistance67 and resistance to diseases (e.g.,
Phakospora pachyrhizi, soybean rust)25,26,68. Glycine
are highly palatable to stock and have been investigated
for their agronomic potential as pasture species, with one
variety of Glycine latifolia commercially released in
Australia69.
Glycine species occur across Australia, with four species
widely distributed: Glycine canescens (silky glycine),
Glycine tabacina (variable glycine), Glycine clandestina
(twining glycine) and Glycine tomentella (rusty glycine)
(Fig. 3b and c). G. canescens had the most desirable distribution as it occurred across the targeted agro-climatic
zones, in particular, within the arid interior (Fig. 3b).
Young plants of G. canescens have been reported to have
particularly good adaptation to low-P stress, partly due to a
high seed P concentration70. G. tomentella occurs within
targeted regions, mainly in the subtropical, subhumid
climatic zone (i.e., E4), but its distribution indicates a
tendency toward more tropical adaptation, and hence was
considered less suitable (Fig. 3b). G. tabacina and
G. clandestina occur predominantly in regions of eastern
Australia with wetter climates and were found to a lesser
extent within the target regions (Fig. 3b and c). Other
evidence suggests that G. tabacina is better adapted to drier
environments than G. tomentella due to its smaller leaflet
size, the exhibition of paraheliotropism and its ability to
maintain photosynthetic gas exchange and chlorophyll
fluorescence at low water availability66. Within their distribution, G. canescens and G. tomentella are commonly found
on sandy soils, G. clandestina on sandy red earths and
G. tabacina is more suited to heavier and deeper soils45.
Less widely distributed Glycine species that occur within
the target regions include G. latifolia (subtropical regions)
and Glycine rubignosa, while Glycine latrobeana was not
suitable as its distribution is limited to cooler, moist
environments of southeastern Australia (Fig. 3c).
All Australian Glycine are perennial twining herbs. Most
are active and flower in the warmer months and usually in
the first year after establishment45. G. clandestina flowers
in spring to early summer, G. tabacina in summer and
G. tomentella in spring and autumn. G. canescens flowers
most of the year and is highly indeterminate45. Some
accessions of G. latifolia grown in Queensland are
exceptionally fast to flower, ranging from 13 to 59 days
to first flower in the establishment year69, suggesting that
germplasm adapted to short growing seasons are available.
In G. tomentella, Jones et al.69 found flowering to be day
length sensitive, with flowering inhibited at longer day
lengths (>16 h), but flowering was initiated with shorter day
lengths (< 12 h). However, variability between accessions
was found. Phenology of node appearance and flowering in
Potential of native Australian legumes as grain crops
G. tomentella is also driven by thermal time, with flowering
occurring after 60 days under warmer conditions (28/24 C
day/night temperatures, i.e., 1560 degree days) and 75 days
under cooler temperatures (24/20 C day/night, i.e., 1650
degree days)71.
Seeds of wild Glycine species are typically moderate
in size (5–10 mg). Measured seed weights are often
between 4 and 6 mg for G. canescens, G. clandestina and
G. tomentella (Table 1). G. latifolia has larger seeds (6.6–
12.5 mg) (Table 1), with seed size of the released forage
cultivar Capella being 12.5 mg69. Glycine seeds are oblong
or ovoid, vary from smooth shiny to roughened dull seed
coats and differ in color between species (G. canescens—
olive-brown, G. clandestina—red–brown, G. tabacina and
G. tomentella—purplish–black).
Surprisingly, no data on the seed composition of wild
Australian native Glycine were found in the literature.
Like soybean and G. soja, Australian native Glycine are
known to produce isoflavones, although these have not been
specifically measured in seeds. Taiwanese wild Glycine
species including G. tabacina and G. tomentella were
reported to have lower isoflavone concentrations in seeds
compared to stems and roots72. Many of these are phytoestrogens have a range of health benefits and applications37.
G. canescens and G. latifolia contain genistin, daidzein and
coumestrol; G. tabacina contains quercetin and kaempterol;
and apigenin was found in G. tomentella, G. tabacina
and G. falcata73. Alkaloids have been reported in Glycine
sericea74, but these are generally not considered to be a
problem in Glycine.
Overall, Australian native Glycine are of significant
interest for further appraisal as a grain crop. They have
attractive seeds of moderate size which potentially contain
chemicals with pharmaceutical applications. The major
constraint for most wild Glycine is their twining/trailing
habit, which is not desirable in a crop plant. Of the Glycine
species, we judge that G. canescens is the highest priority
for further investigation because of its distribution in arid
regions of Australia (Table 3). G. latifolia, because of its
larger seeds and evidence of germplasm with quick maturity, also has a number of suitable attributes. More
information is required on the seed chemistry and seed
yield potential of many species of native Australian
Glycine. Because of their close relationship and potential
for hybridization with soybean, this information would also
be useful for identifying novel or advantageous traits for
soybean breeding.
Glycyrrhiza
Glycyrrhiza is a genus of about 18 species, which includes
only one species native to Australia, Glycyrrhiza acanthocarpa (native liquorice and native lucerne). The genus is
best known for liquorice, which is the product of the roots
of Glycyrrhiza glabra, a species native to the Mediterranean region. Russian liquorice (Glycyrrhiza echinata)
and Chinese liquorice (Glycyrrhiza uralensis) are also
81
cultivated, the latter being important in traditional Chinese
medicine.
G. acanthocarpa occurs from the semi-arid to arid fringe
of southern Australia’s cropping regions, thus appears well
adapted to water-limited environments with a winterdominant growing season (Fig 4a). It occurs in various
habitats and soil types from sandy to clay soils, but is
especially common on soils prone to flooding. It has been
found to be reasonably tolerant of waterlogging and saline
conditions (growth reduced to 59% of control under
120 mM of NaCl solution) (Rogers and Spokes, unpublished data), but performed poorly in a series of field
experiments in waterlogging-prone sites due to poor
establishment and poor biomass production75.
G. acanthocarpa is an erect to semi-prostrate to ascending perennial subshrub growing 1 m high. It flowers
from early spring through to late summer and produces
single-seeded pods covered in hard bristles or prickles.
Advantageously, these pods are indehiscent or tardily
dehiscent, which means pods do not split at maturity, or if
so, quite late46. Seeds of G. acanthocarpa are kidney-shaped
and attractively colored, usually olive-green, mottled with
black46. Seeds are quite small, being about 2.5 mm long and
about 5 mg (Table 1). No documented information was
found on seed yield, seed protein or oil content, or the
presence of bioactive compounds or toxins in seeds. Other
exotic Glycyrrhiza are known to possess a number of medically beneficial properties76; whether these active chemicals occur in the seeds of G. acanthocarpa is unknown.
Overall, we consider G. acanthocarpa to be worth
further appraisal as a grain crop (Table 3). It has a suitable
growth habit, its pods are indehiscent, it has moderate-sized
attractive seeds and its distribution suggests a high suitability to Australia’s more arid cropping regions. More
information is required on the seed chemistry and seed
yield potential of this species.
Hardenbergia
Hardenbergia is a genus of three species, all endemic to
Australia. H. violacea (false sarsparilla, purple coral
tree and happy wanderer) is widely grown as a hardy
ornamental garden plant, with many cultivars available.
H. violacea is a widespread species found in many habitats,
although it generally tends toward higher-rainfall regions
(Fig. 4a). Hardenbergia comptoniana is only found in
southwestern Australia and mainly around the wetter
coastal regions with mean rainfall greater than 700 mm
per annum (Fig. 4a). All species are climbing vines, but
sometimes can assume a subshrub form. Pods are dehiscent
and in some cases these can be explosive77. Hardenbergia
are large-seeded (22–45 mg) (Table 1) and can contain
favorable concentrations of crude protein and oils16
(Table 2). Despite these positive attributes, we consider
Hardenbergia to be of marginal interest as they are primarily adapted to moist environments and their twining
82
L.W. Bell et al.
(a)
growth habit would be a further obstacle to their development as a grain crop.
Indigofera
(b)
(c)
Figure 4. Australian distribution of native Rhyncosia, Glycyrrhiza and Hardenbergia spp. (a), Indigofera spp. (b) and Kennedia
spp. (c) mapped against targeted agro-climatic regions; semi-arid
cropping zone (E2, E3 and E4)—light gray; and the arid interior
(E6 and G)—dark gray (see Fig. 1). Data sourced from Australian
Virtual Herbarium28.
Indigofera is a large genus of about 700 species of which 33
are native to Australia43. They occur throughout the tropical
and subtropical regions of the world, with a few species
reaching the temperate zone. These are mostly shrubs,
although some are herbaceous, and a few can become small
trees up to 5–6 m tall. Most species are dry-season or winter
deciduous. Several of the exotic species (especially
Indigofera tinctoria and Indigofera suffruticosa) are
commercially grown to produce the dye, indigo.
Most Australian Indigofera occur principally in the
tropics and only Indigofera australis (Austral indigo),
Indigofera brevidens (desert indigo), Indigofera colutea
(rusty indigo) and Indigofera linnaei (Birdsville indigo)
occur within our target region to any significant extent
(Fig. 4b). I. australis occurs further south than the other
three species and is found throughout the winter-dominant
rainfall regions (Fig. 4b). I. australis and I. brevidens prefer
sandy soils from granite or sandstone origin and commonly
occur on granite plains and outcrops and river flats78.
These four Indigofera species all have favorable growth
habits and winter–spring growth patterns. I. australis is a
highly variable species, but is often an erect spreading
shrub with flexible stems growing up to 2.5 m tall46. It
flowers in winter to early spring and flowers and pods are
held in leaf axils, distributed along the stem. I. brevidens
and I. colutea are both smaller perennial subshrubs growing
0.4–1 m high, although I. brevidens is often spiny which
may limit its suitability for agriculture46. I. colutea flowers
in autumn, while I. brevidens flowers from spring to early
summer45.
Indigofera seed are small with a squarish, blunt shape
and the seed coat is often spotted. Seeds of I. australis
appear to be larger (about 5 mg) than those of I. colutea and
I. linnaei (< 2 mg) (Table 1). I. australis seeds have been
found to contain 19% crude protein and 2.8% oils, which
was lower than other native legumes tested16 (Table 2).
Indigofera are also known to contain a variety of antinutritional or bioactive compounds such as indospicine and
3-nitropropanoic acid39,78. I. linnaei is known to contain
indospicine in its leaves and seeds which can cause a toxic
condition in horses but not cattle79. I. australis can also
contain HCN and is suspected of being toxic to grazing
livestock while flowering45. Despite the presence of antinutritional factors, a variety of Indigofera species did not
reduce the growth rate of rats fed their seed or leaves78,
indicating their potential as animal feed. Some exotic
species of Indigofera actually have analgesic properties and
have been used historically as anti-inflammatories and for
pain alleviation (e.g., Indigofera articulata, Indigofera
oblongifolia, I. suffruticosa and Indigofera aspalthoides)80.
Whether these qualities are present in Australian Indigofera
Potential of native Australian legumes as grain crops
or if the active compounds are present in the seeds is
unknown.
Of Australia’s Indigofera species, I. australis appears to
have the greatest potential for temperate agriculture due to
its larger seeds and more southern distribution, although it
has been found to have lower protein and fat content than
some other native legumes (Table 3). Seeds of Indigofera
may also offer some novel medicinal uses, although their
chemistry still remains to be explored.
Kennedia
Kennedia have long been identified as legumes with
agricultural potential8. In particular, Kennedia from lowrainfall wheatbelt areas of Australia have been suggested as
possible forage plants (e.g., Kennedia prostrata, Kennedia
stirlingii and Kennedia prorepens)81–83. Yet, no species
have been domesticated, although a number of them are
grown as ornamentals.
The Kennedia genus contains 15 species all of which are
endemic to Australia43. The most widely distributed species
are K. prostrata, found across southern Australia but
mainly in moister regions, and K. prorepens, found throughout the arid regions of central Australia (Fig. 4c). There are
nine Kennedia species endemic to southwestern Australia,
eight of these species have quite localized distributions
mainly along the southern coast or higher rainfall coastal
regions outside the target region (i.e., K. nigricans,
Kennedia glabrata, Kennedia beckxiana, Kennedia carinata, Kennedia eximia, Kennedia stirlingii, Kennedia
macrophylla and Kennedia microphylla) (not shown). The
one more widely distributed western Australian species,
Kennedia coccinea, is predominantly found in the highrainfall regions, although it occurs to a lesser extent in the
target regions (Fig. 4c). Silsbury and Brittan84 observed that
the distribution of K. carinata corresponded to regions with
a >7-month growing season and K. coccinea to regions
with a 6-month growing season, while K. prostrata was
found in drier regions with a shorter growing season
(5 months). There are three Kennedia species only found
in eastern Australia; Kennedia rubicunda has a wide distribution but mainly occurs in higher-rainfall environments
along the east coast (Fig. 4c); Kennedia procurrens is found
within the target region, although almost entirely within the
subtropical subhumid region (E4) (Fig. 4c); and Kennedia
retrorsa has a small distribution outside the target region
(not shown). Kennedia species are mainly found in
woodland or forest habitats and have a preference for light,
well-drained soils. This adaptation to light-textured soils
also suggests that they have some tolerance of drought and
poor soil fertility. Two recent studies show that K. prostrata
and K. prorepens seedlings grew better than some other
perennial legumes under low phosphorus stress, partly due
to high seed phosphorus concentrations70. However, these
same studies show that these two species are particularly
intolerant of high mineral soil phosphorus concentrations
and thus would be suited only to low input agriculture on
83
poor soils. A further problem with Kennedia is a high
degree of seed dormancy which has proved difficult to
overcome.
All Kennedia are evergreen prostrate or climbing
perennials. They are herbaceous but often have woody
stems at their base. Most species display indeterminate
flowering from late winter into early summer with pod
maturity reached about 1 month later. K. prorepens flower
throughout winter beginning in autumn until late spring.
Flowers are open pollinated by insects or birds. Elongated
pea-like pods contain 4–50 seeds. Mature pods are dehiscent, but valves do not twist at maturity. One study has
reported seed production of 200 kg ha - 1 from K. prostrata
at the onset of November at Merredin in the Western
Australian wheatbelt82. This was about 10% of the total
shoot biomass at this time. However, flowering and seed
production did not occur until the second growing season
for K. prostrata82. This is commonly recognized in K.
prostrata, while other Kennedia species (e.g., K. prorepens
and K. coccinea) flower in their first year.
Kennedia have quite large seeds compared to many of
the other native legumes, with many species having seeds
> 10 mg. Seeds up to 44 mg have been measured in
K. prostrata, but the seed size in this and other species
seems to be highly variable (Table 1). The chemical
composition of seeds of some Kennedia species has shown
them to possess high levels of protein and favorable fatty
acids. K. coccinea and K. nigricans were found to have
> 24% protein and fatty acids consisted of 20% saturated,
30% monounsaturated and 50% polyunsaturated fats.
K. nigricans (9%) had higher total fat/oil content than in
K. coccinea (3%)16 (Table 2). K. prostrata seeds have also
been found to contain >22% protein (N%r5.7), which was
concentrated in the embryo and cotyledon85. The embryo
and cotyledons made up only 23.7% of the seed weight
compared to the testa (seed coat) which made up 75% of
the seed weight and contained over 30% of its N and P86.
This contrasts strongly with many domesticated grain
legumes, where the testa consists of a small proportion
of the seed’s dry matter (e.g., Pisum 10.4% and Lupinus
12.7–33.7%) and contains < 5% of the seed’s N and P85,87.
Hence, it appears that significant gains could be made in
improving the total protein yield from Kennedia seed by
selecting for thinner seed coat.
The thick testa in K. prostrata probably imparts the
dormancy and longevity required for seeds to persist over
many years. High levels of seed dormancy have also been
seen in other Kennedia (over 95% seeds were dormant at
maturity in K. rubicunda), which can cause problems for
uniform and reliable germination that would be required in
a crop. Selection for soft, thin seed coats in cultivated grain
legumes has removed the dormancy imparted by a thick
testa, and has enabled nutrients which might have otherwise
gone to this structure to be directed to the embryo31.
Although no major toxicity problems have been documented with Kennedia, Rivett et al.16 found K. nigricans
and K. coccinea seeds to contain significant concentrations
84
of canavanine, 8.1 and 6.0 mol%, respectively. However,
the presence of canavanine in some of these seeds should
not prove an obstacle to their food use since the apparent
toxicity of this compound is low.
Overall, Kennedia are an interesting genus to consider
further as a grain crop. They have large seeds (up to 45 mg)
with advantageous nutritional qualities, some species
produce copious seeds in the first year. As with Glycine,
a major constraint is their twining/trailing habit which is
not favorable in a crop plant. Of the Kennedia species,
K. prorepens has the most desirable distribution and appears well adapted to arid environments and hence was
prioritized for further investigation (Table 3). Germplasm
of K. prostrata also has some desirable adaptations to
challenging environments, but its inability to flower and set
seed in its first year is a major constraint (Table 3).
Lotus
The Lotus genus (bird’s foot trefoils and deer vetches) includes between 70 and 150 species (depending on author).
Several species are cultivated as forage plants in many
regions of the world, but not as a grain crops (e.g., Lotus
corniculatus, Lotus pedunculatus and Lotus glaber). The
genus is large, but only two species are native to Australia,
Lotus australis and Lotus cruentus. Both L. australis and
L. cruentus are found in the target region throughout
southern and inland Australia, but of the two, L. cruentus
may have greater adaptation to the lower-rainfall regions
(Fig. 5a). Both species are found on a wide range of soil
types and habitats and are considered to be drought resistant45.
Both Australian Lotus species can perenniate and flower
in their first year, although L. cruentus often acts as an
annual and produces copious seeds88. L. australis has an
erect-ascending habit growing up to 60 cm in height, while
L. cruentus is more prostrate to ascending46. Flowering can
occur all year round, but mainly occurs in spring with
maturity in early to mid summer46. Plants are pollinated by
insects, commonly bees, although L. cruentus appears to be
reasonably self-compatible (Richard Bennett, unpublished
data).
One major limitation of Lotus species for grain production is the loss of seed due to their continuous flowering
and high propensity for the pod to shatter at maturity. Seed
shattering (dehiscence) is a major problem for seed production in domesticated Lotus species used as forage plants
and seed losses can vary between 5 and 88%33. Lotus
species are generally small seeded, with seeds weighing
between 1.3 and 2.7 g (Table 1). Seed are often smooth,
very round and brown colored with a mottled appearance.
The content of protein, oils or other compounds in the seed
from Lotus species is unknown. Both Australian Lotus
species contain HCN in their shoots, which is associated
with numerous cases of poisoning in cattle and sheep39,45.
However, significant variability in HCN content has been
identified in L. australis, enabling genotypes with low HCN
L.W. Bell et al.
(a)
(b)
(c)
Figure 5. Distribution of Australian native Lotus and Trigonella
spp. (a), a selection of widely distributed Swainsona spp. (b) and
native Vigna spp. (c) mapped against targeted agro-climatic
regions; semi-arid cropping zone (E2, E3 and E4)—light gray; and
the arid interior (E6 and G)—dark gray (see Fig. 1). Data sourced
from Australian Virtual Herbarium28.
Potential of native Australian legumes as grain crops
to be selected and bred89. Highest concentrations of HCN
are found in the leaves and flowers; concentrations are
much lower in the seeds and pods and decrease as pods
mature90. Thus, HCN is unlikely to be a major difficulty in
seeds of Lotus species and could actually be advantageous
in vegetative parts to provide protection from insect
herbivory, while the seed remains palatable.
Australian Lotus species have attracted interest as
potential forage legumes for low-rainfall environments89,
but they seem less well suited as grain crops. They have
quite small seeds, and their unappealing appearance would
limit their novelty as a whole-seed product. Their propensity to shatter is a major agronomic problem and the
lack of interest in using more domesticated Lotus species
for grain production indicates that these species have
limited suitability as grain crops (Table 3).
Rhynchosia
Rhynchosia includes more than 200, mostly tropical, species,
with six species native to Australia. Several species in the
genus are commonly called rosary bean because of their
attractive red, blue, black, mottled or bicolored seeds. The
pantropical species Rhynchosia minima is highly variable
and has four varieties described in Australia [var. amaliae,
australis ( = eurycarpa), minima and tomentosa]. R. minima
has previously been investigated as a potential forage plant
and with many ecotypes that vary in their adaptation and
growth characteristics there is significant opportunity to
exploit this species91.
Most Australian Rhynchosia are restricted to subtropical
and tropical regions of Australia, but the most widespread
species, R. minima (snout bean), is also found across the
arid regions of central Australia and commonly within the
target region (Fig. 4a). R. minima is found in a variety
of habitats but most often on self-mulching heavy clay
soils46,91. However, it has been collected from sands and
sandy loams91. It is regarded as a hardy plant and tolerant
of drought.
R. minima is a slender climbing or trailing perennial
herb. It germinates on summer rains and flowers during
spring–summer and produces abundance of seed46,91. Days
for flowering vary from 43 to 14291. Pod indehiscence was
found in most accessions of Rhynchosia, but some
accessions do retain seeds longer than others91.
Seeds of R. minima are reasonably large, kidney-shaped
and grayish, brown or black and often mottled. The seed
size may vary substantially, ranging from 8.8 to 20.4 mg91.
There is no information on the concentrations of protein or
oils in R. minima seed, but they have been found to contain
some chemicals of pharmaceutical interest including
prodelphinidin (antibiotic), gallic and protocatechuic acid
(antiasthmatic and antioxidant)6,80.
Overall, R. minima warrants further appraisal as a grain
crop (Table 3). It has large attractive seeds and is regarded
as a productive seed producer. Significant variability exists
in important agronomic attributes such as days to flowering,
85
pod indehiscence and seed size, providing the potential to
identify and select desirable genotypes to improve grain
production. Its distribution suggests that it is tolerant of
water-limited environments, but its preference for heavy
clay soils restricts its applicability in some regions.
Swainsona
Swainsona includes 85 species of which 84 are endemic to
Australia. The best known of these species is Swainsona
formosa (Sturt’s desert pea), which is grown as an ornamental flowering plant, but little is known about most of
these species. Swainsona are generally found throughout
the arid interior of Australia with most occurring within
our target region. Many species are not widely distributed,
but the more widely distributed species include S. formosa,
Swainsona canescens, Swainsona colutoides and Swainsona swainsonoides (Fig. 5b). Many species also exhibit
characteristics of plants adapted to dry environments such
as hairy leaves and branches, and a deep tap root.
Swainsona includes plants with annual, biennial and
perennial life cycles and most could be described as small
subshrubs that range from prostrate to semierect92. At least
a few species are winter growing which flower and set seed
in spring (e.g., S. canescens). Most species seem to be
predominantly open pollinated by insects or birds92, but
some have a degree of self-compatibility. A few species
are known to exhibit exceptional fecundity, for example,
well-grown plants of S. canescens are capable of setting
approximately 80,000 seeds (Bennett, unpublished data).
Generally, the genus is described as dehiscent, but a
number of species are known to be indehiscent or tardily
indehiscent (e.g., S. canescens, S. colutoides, Swainsona
pyrophila and Swainsona fraseri), where pods senesce with
seeds are still enclosed. Seeds of Swainsona are usually
small (< 3 mg) and kidney-shaped. However, S. swainsonoides appears to have larger seeds (> 7 mg) than other
species (Table 1). Seeds usually have hard seed coats,
which induce dormancy.
Aborigines ate at least one Swainsona species, Swainsona galegafolia (Darling pea), which was eaten green and
has a similar taste to the common garden pea93. Its green
seed (69% moisture content) contains 31% protein, 33%
carbohydrate, 26% fiber and 6% fat (on a dry matter
basis)93. However, while 64% of the seed is edible, no information about the inedible component was provided. No
information on the constituents of seed of other Swainsona
was found.
Swainsona also gives its name to the toxic alkaloid
swainsonine39; while this is poisonous to livestock, its
effect on humans is unknown. Aplin and Cannon74 report
that the concentration of alkaloids in general (not only
swainsonine) in the vegetative material of other Swainsona
species was high in Swainsona rostellata, moderate
in Swainsona campestris, S. canescens, Swainsona incei,
Swainsona stipularis and low in Swainsona cyclocarpa,
Swainsona flavicarinata and Swainsona occidentalis.
86
Species reputedly or proven to be poisonous when grazed
by livestock include Swainsona galegifolia, Swainsona
sejuncta, Swainsona greyana, Swainsona lessertiifolia,
Swainsona luteola, Swainsona microphylla, Swainsona
oroboides and Swainsona procumbens94. Seeds of S. galegifolia and S. sejuncta contain 2900 and 1700 mg kg - 1 of
swainsonine, respectively27. There are few data on seed
swainsonine concentration in other Swainsona species, but
it would be expected to be negligible in species with low
concentrations in vegetative material. For example, the
swainsonine concentration in the stems or leaves of
S. galegifolia (up to 7500 mg kg - 1) and S. sejuncta (up to
5200 mg kg - 1) is approximately 2.5 times the concentration in the seeds27. Thus species such as S. formosa, which
have low concentrations of swainsonine in leaves (70 mg
kg - 1) and flowers (210–490 mg kg - 1), may have very low
concentrations in seeds27.
Despite the lack of information on many Swainsona
species, a number of them have characteristics which
suggest that they are worthy of further investigation for
their grain production potential (Table 3). In particular,
S. canescens and S. colutoides are high-seed-producing
species, which have delayed dehiscence, an erect growth
habit and are not reported to contain high concentrations of
swainsonine. Being an annual species with low risk of
swainsonine problems, S. formosa may also be of further
interest. Some other widely distributed Swainsona species
(S. swainsonoides, Swainsona purpurea and Swainsona
kingii) may also have potential, but lack information.
Trigonella
Of the 80 species in the genus Trigonella, Trigonella
suavissima (sweet fenugreek) is the only native of
Australia43. This species has been investigated for its
potential as a forage plant95. Several exotic Trigonella
species are important for culinary, nutritional or medical
reasons96. The most widely used is fenugreek (Trigonella
feonum-graecum), which is cultivated throughout semi-arid
regions of the world as an alternative multipurpose crop
that can be grown for grain, forage or green manure97,98.
T. suavissima is a winter-growing annual or ephemeral,
flowering between autumn and spring46. It occurs throughout inland arid environments in central Australia, where
it is typically found on heavy clay soils of river banks,
floodplains and depressions45 (Fig. 5a). It is rarely found on
sandy soils45. It is particularly prevalent in inland Australia
after winter–spring rains or cool-season floods, forming
dense swards on flood plains. Thus, while it occurs in arid
environments its ephemeral life cycle allows it to avoid
severe water stress rather than tolerate water deficit.
T. suavissima has also shown good tolerance of salinity
compared to other native and exotic legumes, with a growth
of 106% of control at 40 mM NaCl concentration99.
T. sauvissima has a desirable growth habit, being
decumbent to ascending and reaching 50 cm in height46.
Few agronomic data are available on the seed production
L.W. Bell et al.
potential of T. sauvissima, yet it is reputed to have a high
level of fecundity45. Collected accessions of the species
have flowered between 111 and 118 days after sowing, but
because of its ephemeral life cyle in arid climates, it is
likely that earlier flowering material exists. It is commonly pollinated by insects, but its self-compatibility is
unknown100. A favorable attribute of T. sauvissima is that it
can be indehiscent or tardily dehiscent, yet fruits are often
shed from plants at maturity92,100.
The seed of one tested accession of T. suavissima is
small (1 mg, Table 1), substantially smaller than its grain
legume relative, fenugreek (9–22 mg)97. The small seeds of
T. suavissima may limit its yield potential as a grain crop.
However, it could have been used as a multipurpose pasture
and crop species (as for fenugreek), as it is regarded as a
valuable and nutritious fodder source where it grows
naturally95. There are no published studies of seed chemical
composition or the presence of bioactive compounds in
T. suavissima. Fenugreek contains a number of bioactive
chemicals and has beneficial medicinal and nutritional
qualities101. The presence of these qualities in T. suavissima
is worth exploring.
Overall, T. suavissima is deserving of further appraisal as
a grain crop (Table 3). It has a suitable growth habit and a
number of desirable agronomic attributes, but in particular
its distribution in arid environments suggests an ability to
avoid or tolerate water stress. Its seeds potentially contain
chemicals with pharmaceutical applications. The major
limitation appears to be small seed size, warranting exploration for germplasm with greater seed size. More
information is required on the seed chemistry and seed
yield potential of this species.
Vigna
The Vigna genus contains a number of species that are
widely grown as grain legumes throughout the world (e.g.,
mung bean (Vigna radiata), azuki bean (Vigna angularis)
and cowpea (Vigna unguiculata)) and some secondary
grain legumes (e.g., Vigna acontifolia (moth bean), Vigna
lanceolata (pencil yam), Vigna mungo (urad bean, black
gram), Vigna subterranea (Bambara groundnut), Vigna
umbellata (rice bean) and Vigna vexillata (zombi pea))98.
Five species of Vigna are indigenous to Australia and one is
endemic (V. lanceolata)102. Vigna radiata ssp. sublobata
is the putative progenitor to the cultivated mungbean
(V. radiata) and is a native of Australia43.
Australian Vigna species are predominately tropical
species or are mainly found in higher-rainfall environments
(Fig. 5c). V. lanceolata is the only species that occurs to
any extent in our target region. It is a highly diverse species
with a number of genotypes that exhibit significant
variation in important agronomic traits (e.g., seed yield,
days to flowering and frost tolerance)103. The key differences in agronomic traits between native Vigna and modern
cultivars are longer time to flowering and maturity, smaller
seed size, higher levels of hardseededness, a more prostrate
Potential of native Australian legumes as grain crops
and twining habit and a lower seed yield and harvest
index104–106. Overall, Australian Vigna are of secondary
interest to us because of their predominantly tropical
distribution and others have previously investigated their
agronomic potential104–106.
Conclusion
Australia has a diverse flora of herbaceous legumes and
their agricultural potential and, especially, their potential to
produce grain products has been little assessed. For many
species, data are sparse and must also be considered with
caution because of the likelihood that past studies have
not adequately captured species’ variability. Nevertheless,
many species possess characteristics that would be useful in
marginal grain growing environments due to their adaptation to arid and semi-arid climates and, sometimes, infertile
soils. A major challenge for a number of species (e.g.,
Glycine, Kennedia and Rhynchosia) is their twining growth
habit. However, this was the case in many of the progenitors of modern legume grain crops (e.g., Glycine,
Vigna, Phaseolus, Pisum and Arachis)31. Similarly, substantial increases in seed size and the removal of seed
dormancy mechanisms have previously been achieved
through plant breeding, and so while many Australian
native species have small seeds, there is a potential to increase seed size. Similarly, seeds of many undomesticated
legumes are likely to have a high proportion of seed coat, as
seen in K. prostrata86, which if reduced could increase
protein yield and reduce problems with hardseededness.
Little information exists on the chemical constituents of
many native Australian legumes, but some have a potential
market because they possess attractive seeds (especially
small-seeded species, e.g., Glycine and Glycyrrhiza) or
because they possess bioactive compounds with prospects
for use as natural medicines (e.g., Glycine, Cullen,
Trigonella and Indigofera). While the germplasm of native
legumes is currently stored in Australian Genetic Resource
Centres107, it is likely that these collections do not come
close to fully representing the diversity present in natural
populations and any serious attempt at domestication of
most native legumes would need to commence with a
comprehensive collection of wild germplasm51.
This paper identifies a number of species with the
greatest immediate potential to be developed as alternative
grain legumes; however, additional basic information on
the seed constituents, phenology, breeding system and reproductive potential of these species is required to narrow
the list further. This is especially necessary in genera where
little current data exist (e.g., Swainsona, Glycyrrhiza and
Crotalaria). This work has been initiated and some preliminary studies will be reported in a forthcoming paper.
Once two or three most promising species are identified,
a preliminary selection and breeding program could
commence to begin the domestication process. This process
could commence with a—perhaps more limited—germplasm acquisition program, but before significant gains
87
could be made, more substantial germplasm acquisition and
characterization activities would be needed. Concurrently,
evidence on the potential applications and market niche for
grain from these species would be required. Ongoing
collaboration with the food and/or nutraceutical industry
would help this process. In addition, agronomic and physiological research should focus on confirming adaptive
characteristics and agronomic suitability and help to focus
future breeding priorities.
Acknowledgements. We would like to thank the Rural Industries Research and Development Corporation (RIRDC) for their
funding of this research. We are also grateful to Jens Berger
and Jon Clements for their comments on the paper.
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