Native Legumes as a Grain Crop for Diversification in Australia

RIRDC Publication No. 10/223

RIRDCInnovation for rural Australia

Native Legumes as a Grain Crop for Diversification in Australia

by Megan Ryan, Lindsay Bell, Richard Bennett, Margaret Collins and Heather Clarke

October 2011

RIRDC Publication No. 10/223 RIRDC Project No. PRJ-000356

© 2011 Rural Industries Research and Development Corporation. All rights reserved.

ISBN 978-1-74254-188-4 ISSN 1440-6845

Native Legumes as a Grain Crop for Diversification in Australia Publication No. 10/223 Project No. PRJ-000356

The information contained in this publication is intended for general use to assist public knowledge and discussion and to help improve the development of sustainable regions. You must not rely on any information contained in this publication without taking specialist advice relevant to your particular circumstances.

While reasonable care has been taken in preparing this publication to ensure that information is true and correct, the Commonwealth of Australia gives no assurance as to the accuracy of any information in this publication.

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Researcher Contact Details

Dr Megan Ryan School of Plant Biology M081 The University of Western Australia 35 Stirling Highway CRAWLEY WA 6009

Phone: 08 6488 2208 Fax: 08 6488 1002 Email: [email protected]

In submitting this report, the researcher has agreed to RIRDC publishing this material in its edited form.

RIRDC Contact Details

Rural Industries Research and Development Corporation Level 2, 15 National Circuit BARTON ACT 2600

PO Box 4776 KINGSTON ACT 2604

Alan Davey Senior Research Manager, New Plant Industries

Phone: 02 6271 4100 Fax: 02 6271 4199 Email: [email protected]. Web: http://www.rirdc.gov.au

Electronically published by RIRDC in October 2011 Print-on-demand by Union Offset Printing, Canberra at www.rirdc.gov.au or phone 1300 634 313

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Foreword

Exotic grain legume crops have played an important role in Australian cropping systems, generating export income, fixing nitrogen and providing a disease break for following cereal crops.

This project is the first systematic assessment of the potential for native Australian herbaceous legumes to be developed as grain legume crops for dry environments in the southern Australian grain belt. Native species may have special application in the face of a drying climate and diminishing supplies of phosphorus, as they are characteristically found in regions with poor soils and low, variable rainfall.

The species studied were Glycine canescens, Cullen tenax, Swainsona canescens, S. colutoides, Trigonella suavissima, Kennedia prorepens, Glycyrrhiza acanthocarpa, Crotalaria cunninghamii and Rhynchosia minima.

Results from this project indicate that the native herbaceous legume species studied have some of the characteristics required for domestication. They have similar oil, protein and fibre contents to existing exotic legume crops and have the potential to be developed as commercial grain legume crops in their own right. All of the species studied in this project showed promise.

There is a scarcity of published information on these species, especially those in the genera Swainsona, Glycyrrhiza and Crotalaria, and there is likely to be considerable variability within each species. Further collection and evaluation of germplasm on these species would be advisable before more intensive trials can be contemplated.

This project was funded from RIRDC Core Funds which are provided by the Australian Government.

This report is an addition to RIRDC’s diverse range of over 2000 research publications and it forms part of our New Plant Products R&D program, which aims to facilitate the development of new industries based on plants or plant products that have commercial potential for Australia.

Most of RIRDC’s publications are available for viewing, free downloading or purchasing online at www.rirdc.gov.au. Purchases can also be made by phoning 1300 634 313.

Craig Burns Managing Director Rural Industries Research and Development Corporation

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Acknowledgments

We thank Sabrina Tschirren and Chelsea Fleming for their technical assistance with the glasshouse experiment; our industry partner, George Weston Foods Ltd for analyses of seed quality; and two CSIRO referees for comments on the review paper. Much of the evaluated germplasm was originally collected as part of activities within the CRC for Plant-based Management of Dryland Salinity.

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Contents

Foreword ...... iii

Acknowledgments...... iv

Executive Summary ...... viii

Introduction ...... 1

Objectives ...... 2

Methodology ...... 3

Section 1: Literature review ...... 3 Section 2: Glasshouse trial ...... 3

Section 1. Literature Review ...... 4

Summary ...... 4 Introduction ...... 4 Approach and desirable plant attributes ...... 5 Potential adaptation to arid and semi-arid environments ...... 5 Harvestability ...... 6 Grain size and yield potential ...... 6 Grain chemistry and nutritional qualities ...... 9 Canavalia ...... 9 Crotalaria ...... 9 Cullen ...... 11 Desmodium ...... 19 Glycine ...... 21 Glycyrrhiza ...... 24 Hardenbergia ...... 26 Indigofera ...... 26 Kennedia ...... 28 Lotus ...... 34 Rhynchosia ...... 39 Swainsona ...... 42 Trigonella ...... 47 Vigna ...... 48

Conclusion ...... 49

Section 2. Glasshouse trial ...... 51

Summary ...... 51 Introduction ...... 51 Materials and Methods ...... 52 Results ...... 56 Discussion ...... 58

Implications...... 61

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Recommendations ...... 62

References ...... 63

Appendix 1 - Workshop Agenda ...... 70

Appendix 2 - Workshop summary and recommendations ...... 71

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Tables

Table 1. Seed size of Australian native herbaceous legumes. Means in brackets, where a range of material has been quoted...... 12 Table 2. Chemical composition of seeds of some native Australian legumes (Rivett et al.1983) ...... 13 Table 3. Prioritisation of species for further investigation as grain legume crops ...... 50 Table 4. The native Australian herbaceous legumes used in the study ...... 54 Table 5. Eight plant growth traits measured in the glasshouse experiment ...... 55 Table 6. Shattering characteristics of 10 native herbaceous legumes and two commercial cultivars. Intact pod drop was not recorded as shattering...... 56 Table 7. Nutrient analyses of grain of native Australian herbaceous legumes and commercially available cultivars...... 57

Figures

Figure 1. Agro-climatic regions of Australia (adapted from Hutchinson et al. 2005) ...... 8 Figure 2. Distribution of a selection of widely distributed Australian native (a) Canavalia, (b) 4 native Crotalaria spp. and (c and d) Cullen spp. mapped against targeted agro-climatic regions: semi-arid cropping zone (E2, E3, E4) – light grey, and arid interior (E6, G) – dark grey (see Fig. 1) ...... 10 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, E4) – light grey; and the arid interior (E6, G) – dark grey (see Fig. 1)...... 20 Figure 4. Australian distribution of native (a) Rhyncosia, Glycyrrhiza and Hardenbergia spp., (b) Indigofera spp., and (c) Kennedia spp. mapped against targeted agro- climatic regions; semi-arid cropping zone (E2, E3, E4) – light grey; and the arid interior (E6, G) – dark grey (see Fig. 1) ...... 33 Figure 5. Distribution of Australian native (a) Lotus and Trigonella spp., (b) a selection of widely distributed Swainsona spp., and (c) native Vigna spp. mapped against targeted agro-climatic regions; semi-arid cropping zone (E2, E3, E4) – light grey; and the arid interior (E6, G) – dark grey (see Fig. 1) ...... 35

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Executive Summary

What the report is about

This report is the first comprehensive assessment of the potential for native Australian herbaceous legumes to be developed as grain legume crops for the drier areas of the Australian grain belt. A review of the literature, along with assessment of various sources of unpublished information, was used to prioritise species. Grain quality and plant growth characteristics relevant to domestication potential, especially harvestability, were then assessed in a glasshouse experiment for 17 species of native legume. Three commercial grain legume cultivars were included for comparison.

Who the report is targeted at

This report is targeted at people interested in using our native flora to improve the sustainability of southern Australian cropping systems, especially in the context of a drying climate. The report is targeted Australia-wide at research funding bodies, scientists, plant breeders, innovative growers and members of the food industry.

Background

Australia has a wealth of native herbaceous legumes which are untapped for their agricultural potential. These species may have special application in the face of climate change and diminishing world supplied of phosphorus as they generally are found in regions with poor soils and low, variable annual rainfall. The world is relying on progressively fewer crops for food security. Small advances in traits such as drought tolerance generally come at a large cost for these species. It is possible that legumes native to Australia could allow exploration of traits that allow adaptation to these challenging conditions or even provide new grain legume crops with desirable traits with a minimal amount of selection/breeding. Such species may be perennials, annuals or, indeed, ephemerals.

Aims/objectives

This project aimed to systematically assess the potential of native Australian herbaceous legumes to be developed as grain legume crops for dry environments in the southern Australian grain belt. A thorough review of existing information was used to produce a list of priority species. Preliminary further assessment was then undertaken in a glasshouse experiment. The results of the project are the first step along a path towards development and commercialisation of a native grain legume crop and are an invaluable guide for future research in this area.

Methods used

The published scientific literature was accessed using standard search procedures and information compiled on the genera Canavalia, Crotalaria, Cullen, Desmodium, Glycine, Glycyrrhiza, Hardenbergia, Indigofera, Kennedia, Lotus, Rhynchosia, Swainsona, Trigonella and Vigna (i.e. 242 species native to Australia). Species were evaluated based on the extent that their natural distribution corresponded to arid and semi-arid climatic regions, as well as information on traits related to harvestability (uniformity of ripening, propensity to retain pod, pod shattering and growth habit), grain quality (seed size, chemistry and colour, absence of toxins) and fecundity. A paper for publication in an international referred journal has been written and submitted. A list of priority species was produced using the information from the review, along with unpublished information from field evaluation of native legumes carried out by Steve Hughes (SARDI Genetic Resource Centre, SA) and Richard Snowball (DAFWA Perth, WA) as part of activities of the CRC for Plant- based Management of Dryland Salinity.

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Seventeen species of native legumes and three commercial grain legume cultivars (chickpea cv. Rupali, field pea cv. Kaspa and lupin cv. Mandelup) were then grown in a glasshouse at the University of Western Australia under conditions of no water or nutrient limitation. Grain quality and plant growth characteristics relevant to domestication potential, especially harvestability, were assessed. Seeds were analysed by George Weston Foods Ltd for fat, protein and dietary fibre content.

Results/key findings

The literature review concluded that Glycyrrhiza and Crotalaria species in particular showed many suitable traits for development as grain legume crops, including erect growth habit, low propensity to shatter, flowers and fruits borne on ends of branches, and moderate to large seeds (5 and 38 mg, respectively). Overall, the species considered highest priority for further investigation were Glycine canescens, Cullen tenax, Swainsona canescens, S. colutoides, Trigonella suavissima, Kennedia prorepens, Glycyrrhiza acanthocarpa, Crotalaria cunninghamii and Rhynchosia minima. The Australian species of Vigna, Canavalia and Desmodium have mainly tropical distributions and are therefore poorly suited for drier areas of the southern Australian grain belt. However, it should be noted that published data on seed yield were rare and, for many other traits, information was also limited. In addition, the information presented on individual species must be considered with caution because of the likelihood that past studies have not assessed the variability within each species.

The glasshouse trial found that the native species generally took longer to begin flowering and their duration of flowering was much longer than commercial cultivars, with eight species flowering continuously once flowering commenced. However, the native species may have flowered sooner if water had been limiting. Harvest index, seed weight/plant and seed size were generally much lower for the native species. Days to podding was generally higher for the native species, with the exception of R. minimai, S. colutoides, T. sauvissima, L. croentus and S. kingii. Aboveground vegetative biomass of native species was generally higher than for the commercial cultivars, with the exception of L. cruentus and S. kingii which produced very little vegetative biomass. Nutritive value of seed of natives was generally similar to that of commercial cultivars. Fat content of native seed ranged from 1.9–5.7%, except for much higher contents in the two Cullen species, C. australasicum and C. tenax, at 10.7% and 10.4%, respectively. Protein content was highest for T. sauvissima (33.6%) which was very similar to lupin (33.2%). All other species, with the exception of R. minima (protein content of 18.8%), had protein contents within the range of the three commercial cultivars (20.3–33.2%). Dietary fibre was also within the range of the cultivated varieties (13.4–42.6%), with most native species within the range 18.8–31.7%. The highest dietary fibre content was found in T. sauvissima (38.6%) which is lower than the highest commercial cultivar, lupin (42.6%). These results indicate that the native species studied have some of the characteristics required for domestication and have similar oil, protein and fibre contents to existing grain legume crops. Further evaluation under commercially- relevant field conditions is now required. Overall, it seems the major limits to domestication of native legumes may be lack of determinate flowering, low harvest index and low seed size, all of which have been improved easily in the domestication of other legumes as grain crops (e.g. lupins). The results of this project can now inform the design of future focused research on domestication of native herbaceous legumes as grain legumes.

Implications for relevant stakeholders

The primary implication of this report is that native herbaceous legumes have potential to be developed as grain legume crops for the drier areas of the southern grain belt, but there is no standout species to recommend for immediate domestication. Further research is required and should use the priority species identified in this project as a guide. However, there is a scarcity of published information on many species, especially those in the genera Swainsona, Glycyrrhiza and Crotalaria and a likelihood of high variability within all the native species. Thus, any future research should commence with collection of additional germplasm and the evaluation of this germplasm under glasshouse and field conditions.

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Recommendations

As considerable further research and development is required before release of a commercial native grain legume cultivar, our recommendations are targeted primarily at the Australian research community (funding bodies and researchers). Recommendations are as follows:

• Further prioritisation of the nine priority species identified in this report should be undertaken with a clear idea of the niche for which they are aimed, from both an agricultural systems and a grain marketing perspective. As such, both agronomic trials and consultation with food and/or nutraceutical industries are essential.

• Further prioritisation will be most effective if informed by collection and assessment of a reasonably wide range of germplasm from all prioritised species. It should also be kept in mind that we were not able to reasonably assess all species in each genus due to lack of information and there may be promising species we have not identified as a priority.

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Introduction

Australia has a wealth of native herbaceous legumes which are untapped for their agricultural potential. In the cropping/pasture systems of the southern Australian grain belt, legumes are included both as grain legume crops and as a component of annual and, increasingly, perennial pastures. Grain legumes play an important role for several reasons. In particular, their seeds are high in protein (e.g. pulses such as lupins, chickpea, field pea) or oil (e.g. soybean). Exports of pulses alone had a gross value of $618 million in 2008/09 (ABARE 2010). Grain legumes are also important in farming systems because they provide nitrogen for following non-legume crops, such as wheat, barley and canola. It is widely recognised that the value of nitrogen contributed by grain and pasture legumes will increase dramatically as rising fuel prices lead to higher costs of fertiliser production. Finally, legumes interrupt the build up of diseases and pests when grown in rotation with cereal crops.

Australian native herbaceous legume species may have special application in the face of climate change and diminishing supplies of phosphorus as many are found in regions with poor soils and low and variable annual rainfall. These plants might be extremely valuable in regions and climates where current exotic grain legume cultivars perform poorly. In particular, growers are seeking alternative crops for marginal areas that experience stresses such as drought, high temperatures or salinity. In addition, the world is relying on progressively fewer crops for food security. Small advances in traits such as drought tolerance generally come at a large cost for these species. Legumes native to Australia could allow exploration of traits for adaptation to these challenging conditions or even provide new crops with desirable traits in this regard, with a minimal amount of selection/breeding. Such species may be perennials, annuals or, indeed, ephemerals. In an Australian context, native grain crops avoid the weed risk issues that come with the introduction of a new overseas species. Exploring the wild native flora provides an exciting and substantial opportunity to identify species with potential as alternative grain crops for the future.

Some Australian grasses and legumes, especially those in the genus Cullen, have been investigated as potential pasture or forage species (Britten & De Lacy 1979; Cohen & Wilson 1981; Dear et al. 2007; Gutteridge & Whiteman 1975; Lodge 1996; Millington 1958; Robinson et al. 2007), 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 grain (Lister et al. 1996), but herbaceous species are more suited to modern broadacre farming systems because they can be harvested mechanically and are more easily removed and rotated with other crops. One Australian grass, Microlaena stipoides, has been investigated to a limited extent for grain production (Davies et al. 2005) yet herbaceous legumes have received little attention. Rivett et al. (1983) 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.

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Objectives

1) Assess native herbaceous legumes for their potential as alternative pulse and oilseed grain crops to identify native legumes with potential for domestication and cultivation

2) Provide basic information regarding seed yield, seed size, protein, starch and oil content, and harvestability on a number of legume species

3) Select 20 species for trialling in glasshouse conditions

4) Rank species with potential

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Methodology

Section 1: Literature review The published scientific literature was accessed using standard search procedures and information compiled on the genera Canavalia, Crotalaria, Cullen, Desmodium, Glycine, Glycyrrhiza, Hardenbergia, Indigofera, Kennedia, Lotus, Rhynchosia, Swainsona, Trigonella and Vigna (i.e. 242 species native to Australia). Species were evaluated based on the extent that their natural distribution corresponded to arid and semi-arid climatic regions, as well as information on traits related to harvestability (uniformity of ripening, propensity to retain pod, pod shattering and growth habit), grain quality (seed size, chemistry and colour, absence of toxins) and fecundity. A paper for publication in an international referred journal has been produced and submitted.

Section 2: Glasshouse trial Material and methods for the glasshouse experiment are presented in Section 2.

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Section 1. Literature Review

Summary Many agricultural systems around the world are challenged by declining soil resources, a drying 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 may aid the continued profitability and environmental sustainability of agricultural systems. This section examines herbaceous native Australian legumes for their capacity to be developed as grain crops. The genera considered are Canavalia, Crotalaria, Cullen, Desmodium, Glycine, Glycyrrhiza, Hardenbergia, Indigofera, Kennedia, Lotus, Rhynchosia, Swainsona, Trigonella and Vigna (i.e. 242 species native to Australia). 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 that their natural distribution corresponded to arid and semi-arid climatic regions, as well as existing information on traits related to harvestability (uniformity of ripening, propensity to retain pod, pod shattering and growth habit), grain qualities (seed size, chemistry and colour, 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 in particular showed many suitable traits, including erect growth habit, low propensity to shatter, flowers and fruits borne on ends of branches, and moderate to large seeds (5 and 38 mg, respectively). The species considered highest priority for further investigation were Glycine canescens, C. tenax, S. canescens, S. colutoides, T. suavissima, K. prorepens, Glycyrrhiza acanthocarpa, Crotalaria cunninghamii, and R. minima.

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 future (Brummer 1998). With drying 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 fertiliser use and reduced reliance on pesticides would improve the sustainability of our agricultural systems (Matson et al. 1997). Perennial grain crops could also provide added benefits of protection of soil from erosion, reduced leaching of water and nutrients, and additional forage for livestock (Bell et al. 2008; Cox et al. 2002). Exploring the wild native flora is an exciting and substantial opportunity to identify species with potential as alternative grain crops for the future (Morris 1997).

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 pasture or forage species (Britten and De Lacy 1979; Cohen and Wilson 1981; Dear et al. 2007; Gutteridge and Whiteman 1975; Lodge 1996; Millington 1958; Robinson et al. 2007), but little work has been conducted on their suitability as grain crops. Woody legumes such as Acacia spp. could have some use an alternative sources of grain (Lister et al. 1996), but herbaceous species are more suited to modern broadacre farming systems because they can be harvested mechanically and are more easily removed and rotated with other crops. One Australian grass, Microlaena stipoides, has been investigated to a limited extent for grain production (Davies et al. 2005), yet herbaceous legumes have received little attention. Rivett et al. (1983) examined a number of native Australian plants for their potential as grain crops and found that the legumes Hardenbergia

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violacea, Crotalaria cunninghamii and Kennedia nigricans warranted further examination as they had relatively large seeds with substantial amounts of crude protein and oil.

There are 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) (Buirchell and Sweetingham 2006), Lupinus luteus (yellow lupin) (Berger et al. 2008) and Desmanthus illinoensis (Illinois bundleflower) (Kulakow et al. 1990). Evidence with these species and advances in our understanding of crop domestication and in the technologies associated with crop breeding may allow for rapid advances in the future (Vaughan et al. 2007). However, domestication of Australian legumes may be more difficult, as there is little or no history of pre- domestication. This means that the net may need to be cast wide, as many species are unlikely to possess traits common to domesticated plants (see Fuller 2007). 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 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 food (Brand-Miller and Holt 1998). Hence Australian native herbaceous legumes have not been subjected to the same pre-domestication pressures that have acted upon other species cultivated by ancient peoples or simply present (as weeds) in early agricultural systems (e.g. Casas et al. 2007; Erskine et al. 1994).

We examined 14 genera of Australia’s herbaceous native legumes for their suitability as grain crops and found that at least nine species are worth exploring further. 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 drying in the future.

Approach and desirable plant attributes Amongst Australia’s legumes, there are 14 genera that contain herbaceous species. Information was gathered on three main aspects: potential adaptation to arid and semi-arid environments (typically less than 700 mm mean annual rainfall); traits related to harvestability, grain size and yield potential; and grain chemistry and nutritional qualities (discussed below). Some genera also include currently 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 hybridisation with the cultivated crop, to either improve agronomic traits of a wild species, or transfer desirable characteristics into the cultivated species (e.g. abiotic or biotic stress tolerance) (Cox et al. 2002). Hybridisation of Australian perennial Glycine species with soybean is one such example (Hartman et al. 1992; Singh et al. 1974). 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 paper (e.g. Martyn et al. 2003).

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 Herbarium (Council of Heads of Australian Herbaria 2008) and matched against Australia’s agro-climatic regions (Hutchinson et al. 2005). Species were

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prioritised if their distribution corresponded to the arid interior (G, E6) or semi-arid environments with sufficient capacity for plant growth in winter-spring (E2, E3, E4) (Fig. 1). Species that occur in these agro-climatic regions are likely to possess adaptations to short or erratic growing seasons, and hot and dry climatic conditions, such as physiological drought tolerance mechanisms or growth patterns which enable them to avoid these stresses (e.g. short-growing season, 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) aeras because plant growth is limited during winter-spring due to lack of moisture and cold, respectively. Although cool-season grain crop production is common in agro-climatic regions E1 (wet ‘Mediterranean’) and D5 (cool-season, wet), the target region was restricted to the less favourable climatic regions with a shorter winter-spring growing season (i.e. E2 – dry ‘Mediterranean’, E3 – temperate, sub-humid, and E4 – sub-tropical, sub-humid). Agro-climates F3 and F4 are warm and wet environments and were also excluded, as they have few climatic stresses that reduce plant growth throughout the year. A wider species distribution was also regarded as favourable 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 process (Weeden 2007) and hence were considered important aspects for evaluating 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 favourable 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)(Smartt 1976). Determinant flowering and indehiscent (shattering) pods would be most advantageous. However, pod dehiscence and indeterminacy in flowering are still a common characteristic in domesticated grain and forage legumes (e.g. soybean, Bailey et al. 1997; birds-foot trefoil, Garcia-Diaz and Steiner 2000). Hence, these were not considered to be disqualifying features for further domestication of wild native legumes.

Grain size and yield potential

Legume grain or seed size is obviously an important aspect as it influences 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 conditions (Leishman et al. 2000). Cultivated grain legumes have large seeds compared to their wild relatives and seed size has been increased substantially through active selection, hence is it is likely seed size of wild legumes will be smaller than that of cultivated grain legumes. For example, seed size has increased at least ten-fold in Phaseolus coccineus (French bean) and by at least five-fold in other legume species (Smartt 1976). This is also demonstrated in germplasm of Lupinus angustifolius (narrow-leaf lupin) where seed size varies substantially from 29 to 244 mg, with ‘wild’ types generally smaller seeded (Cowling et al. 1988). Despite the appeal of species with larger seeds, small seed may be equally appealing, especially if it contains 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-fertilise would be highly desirable. Most domesticated grain legumes are self-fertilising with the exception of Phaseolus coccineus, Vicia faba and Cajanus cajan (Smartt 1976). Some difficulties might occur with outcrossing species that require specific pollinators to achieve optimal seed set, while self-fertilising species, not dependent on pollinators, would be less problematic. Annual species may have a greater overall fecundity, because

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their survival relies on producing viable seeds, but perennial species may be equally productive provided they flower and reproduce in their first year (Cox et al. 2002). Many domesticated annual grain legumes have originated from a perennial life form, most likely because of selection pressure for increasing seed yield (Smartt 1976).

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J1

J2

I3

E7

E4 8 E3 F4 E2 E1 D5 F3

B1 B2

Figure 1. Agro-climatic regions of Australia (adapted from Hutchinson et al. 2005). The target climatic regions include arid and semi-arid regions too dry for field crops (i.e. G, E6) and semi-arid cropping zone where moisture is a major growth limitation, with sufficient capacity for growth in winter-spring (i.e. E4, E3, E2).

Grain chemistry and nutritional qualities

Native legumes found to produce seeds with high concentrations of protein and/or oils/fats would clearly be desirable. In addition, those with favourable 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 which can be toxic (e.g. swainsonine, hydrogen cyanide), but some which have pharmaceutical functions or can provide human health benefits at the correct concentrations (e.g. furanocoumarins, phytoestrogens)(Bourgaud et al. 1990; Setchell 1998). Many cultivated grain legumes also possess some anti-nutritional compounds which have been lowered by breeding (e.g. alkaloids in lupins) (Lin et al. 2009).

Canavalia

The genus Canavalia consists of approximately 70 species mostly of tropical origin. Several species are legume grain crops of secondary importance, including common jack-bean (Ca. ensiformis), sword bean (Ca. gladiata) and Ca. cathartica. Raw seeds of Canavalia contain a number of anti- nutritional factors including phenolics, tannins, saponins, concanavalin A, canavanine, cyanogenic glycosides and HCN (Belmar and Morris 1994; Jermyn 1985). Though some of these compounds may have beneficial properties; Canavalia are famous for the presence of the lectin, concanavalin A which has commercial importance as a reagent in glycoprotein biochemistry and immunology (Rüdiger and Gabius 2001; Sridhar and Seena 2006). Four species of Canavalia are found in Australia but none are endemic: Ca. rosea, Ca. carthartica, Ca. sericea and Ca. papuana. These are mostly found in tropical, coastal hinterland regions (Fig. 2a). While Ca. 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 the target climatic regions, they are not considered further here, though they may have some potential as a tropically adapted legume crop. They possess large seeds and are a rich protein source (Jermyn 1985; Sridhar and Seena 2006).

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. Some 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 Australia (Legume Web - ILDIS World Database of Legumes vers. 10 2008). Some exotic species of Crotalaria have agronomic uses (e.g. Cr. spectabilis, Cr. ochroleuca, Cr. longirostrata and Cr. juncea (sunn hemp)) (Legume Web - ILDIS World Database of Legumes vers. 10 2008). The Australian native Crotalaria species are found mainly in tropical regions. Four species occur further south in the target region: Cr. eremaea (desert rattlepod), Cr. mitchelli (yellow rattlepod), Cr. cunninghamii (green birdflower or parrot pea) and Cr. dissitiflora (plains rattlepod) (Fig. 2b). Cr. eremaea and Cr. cunninghamii may be particularly drought tolerant species, occurring mainly on sandy or well-drained soils in low rainfall regions of central Australia (The Australian Arid Lands Botanic Garden 2008). Cr. mitchelli occurs on sandy soils in the tropical and subtropical areas of the east coast with >500 mm MAR (Cunningham et al. 1981). Cr. dissitiflora occurs on heavy clay soils also in the subtropics and tropics although further inland and in lower rainfall regions than Cr. mitchellii (Cunningham et al. 1981).

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(a) (b)

(c) (d)

Figure 2. Distribution of a selection of widely distributed Australian native (a) Canavalia, (b) 4 native Crotalaria spp. and (c and d) Cullen spp. mapped against targeted agro- climatic regions: semi-arid cropping zone (E2, E3, E4) – light grey, and arid interior (E6, G) – dark grey (see Fig. 1). Data sourced from Australian Virtual Herbarium (Council of Heads of Australian Herbaria 2008).

Crotalaria includes annual, biennial and perennial species that 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: Cr. cunninghamii is an erect perennial sub-shrub growing to 1 m or more high, Cr. eremaea is an erect sub-shrub 0.5–1 m high, Cr. dissitiflora is an erect-sprawling short-lived perennial <30 cm high, and Cr. mitchellii is an erect–decumbent woody forb about 60 cm high (The Australian Arid Lands Botanic Garden 2008). All these species flower in winter-spring (Cr. cunninghamii sometimes in autumn) and are generally open pollinated by insects. Cr. dissitiflora has shed its leaves during winter (Cunningham et al. 1981). A notable and advantageous characteristic of these species is that flowers and pods are borne at branch ends (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010), which could favour mechanical harvesting. However, some Crotalaria are elastically dehiscent which may be problematic, while others have passive dehiscence (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010).

Some Crotalaria have large seeds (e.g. 38 mg in Cr. cunninghamii) (Table 1), while others are smaller (e.g. 2–3 mm long in Cr. dissitiflora). Seeds are often smooth and vary in colour (yellow in Cr. dissitiflora, greenish-grey in Cr. mitchellii and red-brown in Cr. smithiana). Due to its large seed

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size and substantial protein and oil content (Table 2), Cr. cunninghamii was identified as a species worthy of further investigation (Rivett et al. 1983). Toxic pyrrolizidine alkaloids are produced by some members of this genus, which can be poisonous to livestock (Everist 1974), but whether these are present in seeds of the four species which occur in the target region is unknown. Cr. dissitiflora is suspected of poisoning livestock but there is conflicting evidence (Cunningham et al. 1981). Everist suggests that the toxicity might be lost when plants are cut (Everist 1974). Cr. eremaea is often eaten by sheep, suggesting low or no alkaloid problems and Cr. cunninghamii is reputedly edible to humans without any indication that prior treatment is necessary (The Australian Arid Lands Botanic Garden 2008; Crib and Crib 1976).

Overall, Australian Crotalaria seem to have a number of characteristics that suggest they could be further investigated for their potential as grain crops. In particular, Cr. cunninghamii has a desirable growth habit, produces large seeds which contain high levels of protein and some oil, and it 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. Cr. eremaea, Cr. mitchelli, and Cr. dissitfolia), but they may also warrant further investigation.

Cullen

The Cullen genus includes 32 species, 25 of which are endemic to Australia (Grimes 1997). Cullen has been explored as forage plants in the past and again recently in Australia (Bennett et al. in press; Burbridge 1980; Dear et al. 2007). 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 species in Australia are 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 mm to 1300 mm (Bennett et al. in press). All species of Cullen occur within the target region and 12 occur mainly in low rainfall environments with an annual average rainfall ≤400 mm (Bennett et al. in press). Of these species C. australasicum, C. graveolens, C. pallidum and C. discolor occurred mainly in the lower rainfall regions (Fig. 2c and 2d). C. 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 plants (Dear et al. 2007; Suriyagoda et al. 2010). All species have a deep tap-root which can become woody in the perennial species. Roots of C. patens (syn. P. eriantha) have been reported to penetrate to a depth of 4 m, and this was associated with the drought resistance of this species (Kerridge and Skerman 1968). Another evident adaptation to drought is the dense coverings of glandular hairs on the leaves of some Cullen species (e.g. C. pallidum, C. patens). Most Cullen species don’t seem to have strong soil type associations and can be found on a range of soil types (e.g. C. australasicum), but some species have particular preferences; for example, C. tenax seems to prefer heavy clay soils and C. pallidum is found predominately on deep sand dunes and sandy soils (Bennett et al. in press; Cunningham et al. 1981; Grimes 1997).

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Table 1. Seed size of Australian native herbaceous legumes. Means in brackets, where a range of material has been quoted. Species Seed mass (mg) Reference Crotalaria cunninghamii 38.5 Rivett et al. 1983 Cullen australasicumA 5.5 Dear et al. 2007 Cullen cinereumA 4.2 Jurado et al. 1991 Cullen cinereumA 4.1 Bourgaud et al. 1990 Cullen cinereumA 5.5 Unpublished data Cullen patensA 8.4 Jurado et al. 1991 Cullen patensA 2.8B Silcock and Smith 1990 Cullen plumosumA 11.3 Bourgaud et al. 1990 Cullen tenaxA 4.4 Unpublished data Glycine canescens 5.6 Jurado et al. 1991 Glycine canescens 5.9–8.9 Unpublished data Glycine clandestine 4.2 Auld and O'Connell 1991 Glycine tomentella 5.0 Silcock and Smith 1990 Glycine latifolia 11.1 McDonald 2002 Glycine latifolia 6.6–12.5 Jones et al. 1996 Glycyrrhiza acanthocarpa 5.4 Unpublished data Hardinbergia violacea 38.5 Rivett et al. 1983 Hardinbergia violacea 22 Auld and O'Connell 1991 Hardinbergia comptoniana 38.3–45.2 Bell et al. 1995 Indigofera colutea 1.1 Jurado et al. 1991 Indigofera linnaei 1.8 Jurado et al. 1991 Indigofera linnaei 1.9 Unpublished data Indigofera australis 5.2 Rivett et al. 1983 Kennedia coccinea 8.8 Rivett et al. 1983 Kennedia coccinea 26.3 Bell et al. 1995 Kennedia eximia 9.0 Unpublished data Kennedia nigrans 15.6 Rivett et al. 1983 Kennedia prorepens 12.4 Denton et al. 2006 Kennedia prorepens 6.6 Unpublished data Kennedia prostrata 29.7 Bell et al. 1995 Kennedia prostrata 31.9 Hocking and Kortt 1987 Kennedia prostrata 44.4 Unpublished data Kennedia prostrata 9.3 Unpublished data Kennedia rubicunda 24.4 Auld and O'Connell 1991 Lotus australis 2.7 Denton et al. 2006 Lotus australis 1.31.9 Unpublished data Lotus cruentis 1.8 Moles et al. 2003 Rhynchosia minima 11.8 Silcock and Smith 1990 Rhynchosia minima 8.8–20.4 (12.1) Harding et al. 1989 Rhynchosia minima 9.4 Jurado et al. 1991 Rhynchosia minima 16.8 Unpublished data Swainsona canescens 2.1 Jurado et al. 1991 Swainsona canescens 1.3 Unpublished data

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Swainsona kingii 2.6 Unpublished data Swainsona purpurea 2.8 Unpublished data Trigonella suavissima 1.0 Unpublished data Vigna radiata ssp. Sublobata 7.4–27.0 (13.4) Lawn and Rebetzke 2006 Vigna lanceolata 20–34 Lawn and Holland 2003 Vigna vexillate 8.1–17.7 James and Lawn 1991 Vigna vexillate 7.0–19.5 (11.1) Grant et al. 2003 AIndicates whole fruit (pod + seed); BImmature seeds harvested

Table 2. Chemical composition of seeds of some native Australian legumes (Rivett et al. 1983). Seeds were obtained from a commercial native seed service and had presumably been collected from the wild. Species Crude Fat Fatty acid composition (% of total fat) A Protein (%) (%) 16:0 18:0 18:1 18:2 18:3 P:S ratioB Crotalaria cunninghamii 23.3 3.8 17.2 6.8 26.0 46.5 3.3 2.1 Hardenbergia violacea 21.0 8.1 12.1 5.1 23.2 55.9 5.0 3.4 Indigofera australis 18.8 2.8 17.6 3.8 26.3 45.6 6.2 2.4 Kennedia coccinea 27.4 3.0 12.3 7.0 29.2 43.9 6.4 2.6 Kennedia nigricans 23.9 9.1 14.9 6.5 30.2 41.0 4.4 2.1 AProtein was calculated as 5.7 x %N; BRatio of polyunsaturated to saturated fatty acids

The Australian species of Cullen include shrubs, sub-shrubs and herbs and a number have a favourable growth habit and phenology. Nineteen species are herbaceous or semi-herbaceous, of which 16 are perennial or short-lived perennials. C. graveolens, C. plumosum and C. walkingtonii are annual or biennial (Grimes 1997). Most Australian taxa will bloom in the first year (Grimes 1997). Flowering mainly occurs in spring, but indeterminate flowering will continue throughout the year provided sufficient moisture is available (Grimes 1997; Kerridge and Skerman 1968). In a glasshouse study, Bourgaud et al. (1990) recorded that flowering occurs around 40 days after germination in C. cinereum (about 900 °d) and around 60 days after germination in C. plumosum (1340 °d). Flowering of C. australasicum and C. patens is controlled by day length according to Britten and De Lacy (1977), with long-day treatments (i.e. less than 12–13 hours dark) inducing flowering. They also found that genotypes vary in their response which might be useful in breeding programs in these species. In C. australasicum, flowering and fruiting times are extremely variable in the first year but with greater synchrony in the second year (Dear et al. 2007).

Many Cullen species seem to be capable of self-pollinating. Britten and Dundas found that erect types in the Psoralea patens complex (i.e. C. australasicum) were 50–75% selfing, while prostrate and semi-erect lines (i.e. C. patens) were outcrossing only (Britten and Dundas 1985). Bougaud et al. (1990) noted that C. cinereum and C. plumosum were capable of self-pollinating. Using microsatellite markers, Kroiss et al. (2009) estimated the outcrossing rate in C. australasicum to be at least 3–13% and hybrids were formed with C. pallidum, but not C. discolour or C. patens.

A couple of studies found Cullen to allocate significant resources to reproduction and produce useful amounts of seed. Bourgaud et al. (1990) found seed yields up to 1.65 g/plant (47% of DM) from C. cinereum and 1.75 g/plant for C. plumosum (60% of DM). The higher yield from C. plumosum was due to the greater seed mass (11.3 mg), while whole plant biomass was less than C. cinereum. Production of seeds from C. tenax has been measured at 22 g/plant (4820 seeds) (Bennett, unpublished data). Kerridge and Skerman recorded that reproductive parts of C. pallidum made up 42% of plant biomass when plants were left to grow for 12 weeks (Kerridge and Skerman 1968). 13

Cullen species are characterised by indehiscent (non-shattering) fruits with the seeds adherent to the pericarp (pod). Fruit sizes of Cullen typically range from 4–6 mg, although 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 Smith (1990), 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 varies. Skerman reported that ripe pods of C. patens drop to the ground and seed harvesting needs to be performed by suction (Skerman 1957). The pods of several species (e.g. C. australasicum, C. patens, C. pallidum, C. discolor) fall from the plant enclosed in the calyx, which can be very hairy (Grimes 1997). This, and the adherence of the seed to its pod, also poses some complications about the ability to thresh seed of Cullen, unless processing used the whole fruit. Dear et al. (2007) state that 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 angelicin (Innocenti et al. 1997a; Innocenti et al. 1997b; Nguyen et al. 1997). Furanocoumarins are potent photosensitising agents that can cause phototoxic reactions but they are also pharmaceutically useful for treatment of skin disorders such as psoriasis, vitiligo, leucoderma and leprosy (Innocenti et al. 1977). The seeds of Australian Cullen species contain between 1000 and 8000 mg/kg DW of furanocoumarins (depending on species) and have been proposed as potential sources for pharmaceutical use (Bourgaud et al. 1990; Innocenti et al. 1977; Nguyen et al. 1997). C. cinereum (syn. Psoralea cinerea) and C. plumosum were identified with the highest levels of furanocoumarins, but they have also been measured in C. lachnostachys (syn. P. lachnostachys) and C. pustulatum (syn. P. pustulata) and are likely to exist in many other species. The fruits generally contain the highest concentration of furanocoumarins, up to 5500 ppm in C. corylifolia (native to India), with most (>70%) found in the cotyledon of the fruit (Innocenti et al. 1997b; Raghav et al. 2003). Vegetative material can also contain significant levels of furanocoumarins (up to 1600 ppm) (Innocenti et al. 1984; Innocenti et al. 1997b), which can 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 also contain the flavonoid, diadzein, which is increasingly studied because of its activity in cancer prevention and treatment (Bouque et al. 1998). Daidzein has been found in the fruits of two Australian species, C. cinereum (8.2 mg/g DW) and C. tenax (27.5 mg/g DW) and was also present in the stems (Bouque et al. 1998; Nguyen et al. 1997). C. patens and C. cinereum also contained lectins and trypsin inhibiting proteins (Jermyn 1985).

A number of Cullen species have invoked significant interest in their potential for domestication for agriculture in the past and again more recently because of their tolerance to arid environments and challenging soils (e.g. P deficient) (Dear et al. 2007; Hayes et al. 2009; Li et al. 2008; Pang et al. 2010). Their ability to produce large amounts of seed, self-compatibility, erect growth habit, ability to grow and flower in the first year (some within 40 days after germination), and moderate seed size in some species are all favourable attributes for development as a grain crop. Another exciting attribute is the likely presence of furanocoumarins which may provide a pharmaceutical market for Cullen seed. However, the problem of separating seed from calyx is a significant issue and their market success would rely on using the whole fruit. A number of Cullen species warrant further investigation for potential as grain crops (Table 3), but high priority species would have a reputation for high seed production (e.g. Cullen tenax, Cullen cinereum) or display an annual life-cycle (e.g. Cullen graveolens) (see also Plates 1–14).

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Plate 1. Cullen australasicum can grow under very hot and dry conditions and is of interest to those developing perennial pastures in Australia

Plate 2. Cullen cinereum can produce seed even in very short seasons and flower in less than 6 weeks if conditions are poor

Plate 3. Each flower on this tightly packed inflorescence of Cullen cinereum results in a pod with a single seed, even in the absence of pollinating vectors

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Plate 4. Pods, each containing a single seed, are packed tightly along the inflorescence of Cullen cinereum

Plate 5. The stature of Cullen cinereum may allow its seed to be easily harvested by conventional headers.

Plate 6. Cullen species are a preferred food plant for some native butterflies such as the Chequered Swallowtail larvae (Papilo demodocus), seen here grazing Cullen cinereum near Carnarvon, Western Australia

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Plate 7. Cullen discolor produces copious seed, however it falls easily from the plant and would probably require harvest by suction machinery

Plate 8. The inflorescences of Cullen discolor are located in the middle of a strongly prostrate canopy making harvest difficult

Plate 9. Cullen leucanthum has an ideal habit and stature for header harvesting

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Plate 10. A close up of Cullen leucanthum flowers with a fly possibly performing pollination duties.

Plate 11. Cullen patens produces copious seed, however it falls easily from the prostrate plants and would probably require harvest by suction machinery

Plate 12. Cullen tenax is perennial, palatable and produces seed in the upper canopy; characteristics that may make it a dual use legume, able to be grazed by livestock and harvested for seed

Plate 13. Cullen cinereum seed

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Plate 14. Cullen tenax seed

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 Australia (Legume Web - ILDIS World Database of Legumes vers. 10 2008). No Desmodium species are grown as grain crops but some are cultivated as forage for livestock (e.g. D. intortum, D. uncinatum) and as living mulch or green manures.

Most Australian Desmodium occurs in the tropics and subtropics and only three species are distributed within the target region: D. varians (slender tick-trefoil), D. campylocaulon (creeping tick- trefoil) and D. brachypodum (large tick-trefoil) (Fig. 3a). D. varians occurs the furthest south into the temperate regions of Australia, but here 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 or twining long-lived perennial sub-shrubs growing to 60–100 cm high which flower from late spring to autumn (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010). 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) (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010). Desmodium pods also don’t split or dehisce at maturity; they have a saw-like pod with segments that separate at maturity enclosing an individual seed (known as a loment) which are dispersed individually (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010). 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 Desmodium species to be of marginal interest because of the little information on the agronomic and seed attributes, and their tendency towards moister and summer-dominant rainfall environments.

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(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, E4) – light grey; and the arid interior (E6, G) – dark grey (see Fig. 1). Data sourced from Australian Virtual Herbarium (Council of Heads of Australian Herbaria 2008).

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Glycine

Australia is the centre for 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 G. soja make up the subgenus Soja which originates in south-east Asia. Numerous attempts to hybridise wild Glycine and G. max have been made and have mainly been successful with tetraploid (2n=80) types of G. tomentella (Cox et al. 2002). Australian Glycine species have been investigated for traits that may be beneficial for soybean improvement, such as drought tolerance (Kao et al. 2003), 2-4D resistance (Hart et al. 1991) and resistance to diseases (e.g. Phakospora pachrhizi, soybean rust) (Burdon 1988; Hartman et al. 1992; Singh et al. 1974). Glycine which are also highly palatable to stock have been investigated for their agronomic potential as pasture species, with one variety of G. latifolia released commercially in Australia (Jones et al. 1996).

Glycine species occur across Australia, with four species widely distributed: G. canescens (silky glycine), G. tabacina (variable glycine), G. clandestina (twining glycine) and G. tomentella (rusty glycine) (Fig. 3b and 3c). 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 are reported to have particularly good adaptation to low-P stress, partly due to a high seed P concentration (Pang et al. 2010). G. tomentella occurs within targeted regions, mainly in the subtropical, sub-humid climatic zone (i.e. E4), but its distribution indicates a tendency towards 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 to a lesser extent within the target regions (Fig. 3b and 3c). Other evidence suggests that G. tabacina is better adapted to drier environments than G. tomentella due to its smaller leaflet size, exhibition of paraheliotropism and its ability to maintain photosynthetic gas exchange and chlorophyll fluorescence at low water availability (Kao et al. 2003). 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 soils (Cunningham et al. 1981). Less widely distributed Glycine species that occur within the target regions include G. latifolia (subtropical regions) and G. rubignosa, while G. latrobeana was not suitable as its distribution is limited to cooler, moist environments of south-eastern 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 establishment (Cunningham et al. 1981). G. clandestina flowers in spring to early summer, G. tabacina in summer (Plate 15), and G. tomentella in spring and autumn. G. canescens flowers most of the year and is highly indeterminate (Cunningham et al. 1981). 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 year (Jones et al. 1996), suggesting that germplasm adapted to short growing seasons may be available. In G. tomentella, Jones et al. (1996) found flowering to be day length sensitive, with flowering inhibited at day lengths >16 hours, but flowers at day lengths <12 hours. However, variability between accessions was observed. Phenology of node appearance and flowering in 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 °days) and 75 days under cooler temperatures (24/20°C day/night, i.e. 1650 °days) (Kenworthy et al. 1989).

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 to12.5 mg) (Table 1), with seed size of the released forage cultivar Capella being 12.5 mg (Jones et al. 1996). Glycine seeds are oblong or ovoid, vary from smooth shiny to roughened dull seed coats and differ in colour 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 was found in the literature. Although, like soybean and G. soja, Australian native Glycine are known to produce isoflavones, although these have not been measured in seeds specifically. Many of these are phytoestrogens and may have a range of health benefits and applications (Setchell 1998). G.

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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. falcata (Vaughan and Hymowitz 1984). Taiwanese wild Glycine species including G. tabacina and G. tomentella were reported to have quite low isoflavone concentrations in seeds compared with stems and roots (Lin et al. 2005). Alkaloids have been reported in G. sericea (Aplin and Cannon 1970), 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 conducive as 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; Plates 19– 20). 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 hybridisation with soybean, this information would also be useful for identifying novel or advantageous traits for soybean breeding.

Plate15. These open and outcrossing chasmogamous flowers of the amphicarpic Glycine tabacina are produced seasonally in conjunction with non-opening cleistogamous flowers that do not require pollinating vectors

Plate 16. The twining habit displayed by these Glycine canescens plants is common among progenitors of many modern crop legumes

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Plate 17. Seed pods of Glycine canescens are produced in the midst of the herbaceous canopy

Plate 18. This Glycine canescens plant is twining through and over a tree and producing copious pods on the underside of the tree canopy

Plate 19. Glycine canescens seed

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Plate 20. Glycine canescens seed (a different accession to Plate 19)

Glycyrrhiza

Glycyrrhiza is a genus of about 18 species which includes only one species native to Australia, Gl. acanthocarpa (native liquorice, native lucerne). The genus is best known for liquorice, which is the product of the roots of Gl. glabra, a species native to the Mediterranean region. Russian liquorice (Gl. echinata) and Chinese liquorice (Gl. uralensis) are also cultivated, the latter being important in traditional Chinese medicine.

Gl. 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 winter-dominant 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 is reasonably tolerant of waterlogging and saline conditions (growth reduced to 59% of control under 120 Mm NaCl solution) (Rogers & Spokes, unpublished data), but performed poorly in a series of field experiments in waterlogging-prone sites due to poor establishment and poor herbage production (Li et al. 2008).

Gl. acanthocarpa is an erect to semi-prostrate to ascending perennial sub-shrub growing to 1 m high. It flowers from early spring through to late summer and produces one-seeded pods covered in hard bristles or prickles. Advantageously, these pods are indehiscent or tardily dehiscent, which means pods don’t split at maturity or if so quite late (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010). Seeds of Gl. acanthocarpa are kidney-shaped and attractively coloured, usually olive-green, mottled with black (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010). 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 properties (Ross 2001); whether these active chemicals occur in the seeds of Gl. acanthocarpa is unknown.

Overall, we consider Gl. acanthocarpa to be worth further appraisal as a grain crop (Table 3; Plates 21–24). 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.

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Plate 21. An inflorescence of Glycyrrhiza acanthocarpa produces a characteristic resinous, sweet smelling substance

Plate 22. Pods of Glycyrrhiza acanthocarpa with rough, hooked projections, each containing a single seed are retained well on plants

Plate 23. The habit of Glycyrrhiza acanthocarpa, growing on the side of Great Eastern Hwy in the wheatbelt of Western Australia: such plants could possibly be header harvested

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Plate 24. Glycyrrhiza acanthocarpa seed

Hardenbergia

Hardenbergia is a genus of three species, all endemic to Australia. H. violacea (False Sarsparilla, Purple Coral Tree, 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 towards higher rainfall regions (Fig. 4a). H. comptoniana is only found in south-western 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 sub-shrub form. Pods are dehiscent and in some cases these can be elastic (Wilson and Wilson 2006). Hardenbergia are large- seeded (22–45 mg) (Table 1) and can contain favourable concentrations of crude protein and oils (Table 2; Rivett et al. 1983). Despite these positive attributes, we consider Hardenbergia to be of marginal interest as they are primarily adapted to moister environments and their twining growth habit and highly dehiscent pods would be major limitations for development as a grain crop.

Indigofera

Indigofera is a large genus of about 700 species and 33 species are native to Australia (Legume Web - ILDIS World Database of Legumes vers. 10 2008). They occur throughout the tropical and subtropical regions of the world, with a few species reaching the temperate zone. The species are mostly shrubs, though some are herbaceous, and a few become small trees up to 5–6 m tall. Most species are dry-season or winter deciduous. Several of the exotic species (especially I. tinctoria and I. suffruticosa) are grown commercially to produce the dye indigo.

Most Australian Indigofera occur principally in the tropics; only I. australis (Austral indigo), I. brevidens (Desert indigo), I. colutea (rusty indigo) and I. 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 flats (Aylward et al. 1987).

These four Indigofera species have favourable 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 tall (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010). It flowers in winter to early spring and flowers and pods are held in leaf axils, distributed along the plants stem. I. brevidens and I. colutea are both smaller perennial sub-shrubs growing 0.4–1 m high, although I. brevidens is often spiny which may limit its suitability for agriculture (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010). I. colutea flowers in autumn while I. brevidens flowers spring to early summer (Cunningham et al. 1981).

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Indigofera seeds 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 contain 19% crude protein and 2.8% oils, which was lower than other native legumes tested (Table 2; Rivett et al. 1983). Indigofera are also known to contain a variety of anti-nutritional or bioactive compounds such as indospicine and 3-nitropropanoic acid (Aylward et al. 1987; Jermyn 1985). I. linnaei is known to contain indospicine in its leaves and seeds which can cause a toxic condition in horses but not cattle (Gracie 1996). I. australis can also contain HCN and is suspected of being toxic to grazing livestock while flowering (Cunningham et al. 1981). Despite the presence of anti-nutritional factors, a variety of Indigofera species did not reduce growth rates in rats fed their seed or leaves (Aylward et al. 1987). This shows that some accessions and species might be identified that don’t contain these toxins and could be still be useful animal feeds. Some exotic species of Indigofera have analgesic properties which have been used historically as anti-inflammatories and for pain alleviation (e.g. I. articulata, I. oblongifolia, I. suffruticosa and I. aspalthoides) (GRIN (Germplam Resources Information Network)). Whether these qualities are present in Australian Indigofera 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 lower protein and fat contents than some other native legumes (Table 3; Plates 25–26). Seed of Indigofera may also offer some novel medicinal uses, although their chemistry remains to be explored.

Plate 25. Copious inflorescences and pods of Indigofera linnaei are produced in the canopy of the strongly prostrate plants

Plate 26. Indigofera linnaei seed

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Kennedia

Kennedia species have long been identified as legumes with agricultural potential (Millington 1958). In particular, Kennedia from low rainfall wheatbelt areas of Australia have been suggested as possible forage plants (e.g. K. prostrata, K. stirlingii and K. prorepens) (Cocks 2001; Silsbury 1952; 1958). Yet, no species have been domesticated, though a number are grown as ornamentals.

The Kennedia genus contains 15 species all of which are endemic to Australia (Legume Web - ILDIS World Database of Legumes vers. 10 2008). The most widely distributed species are K. prostrata, found across southern Australia but mainly in moister regions, and K. prorepens which has a highly desirable distribution as it is found throughout the arid regions of central Australia (Fig. 4c). There are nine Kennedia species endemic to south-western Australia, eight of these species have localised distributions mainly along the southern coast or higher rainfall coastal regions outside the target region (i.e. K. nigricans, K. glabrata, K. beckxiana, K. carinata, K. eximia, K. stirlingii, K. macrophylla and K. microphylla)(not shown). A more widely distributed Western Australian species, K. coccinea, is predominantly found in high rainfall regions, though it occurs to a lesser extent in the target regions (Fig. 4c). Silsbury and Brittan (1955) observed that the distribution of K. carinata corresponded to regions with a greater than seven month growing season, K. coccinea with a six month growing season, whilst K. prostrata was found in drier regions with a shorter growing season (five months) (Silsbury and Brittan 1955). There are three Kennedia species found only in eastern Australia: K. rubicunda has a wide distribution but mainly occurs in higher rainfall environments along the east coast (Fig. 4c), K. procurrens is found within the target region though almost entirely within the subtropical sub-humid region (E4) (Fig. 4c) and K. retrorsa has a small distribution outside the target region (not shown). Kennedia species are mainly found in woodland or forest habitats and prefer light, well-drained soils. This adaptation to light-textured soils suggests that Kennedia is drought tolerant and perhaps infertile. Two recent studies show that K. prostrata and K. prorepens seedlings grew better than some other perennial legumes under low P stress, partly due to high seed P concentrations (Pang et al. 2010). However, these same studies found that these two species are particularly intolerant of high soil P concentrations and thus may be only suited to low input agriculture on 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 one month later (Plates 27–34). K. prorepens may 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 don’t twist at maturity. One study reported seed production of 200 kg/ha from K. prostrata at the start of November at Merredin in the Western Australian wheatbelt (Silsbury 1952). 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. prostrata (Silsbury 1952). This is commonly recognised in K. prostrata, while other Kennedia species (e.g. K. prorepens, K. coccinea) flower in their first year.

Kennedia have large seeds compared to many other native legumes, with many species having seeds >10 mg. Seeds up to 44 mg have been measured in K. prostrata, but seed size in this and other species is highly variable (Table 1; Plates 35–37). Seeds of some Kennedia species have high levels of protein and favourable fatty acids. K. coccinea and K. nigricans have >24% protein, with fatty acids consisting of 20% saturated, 30% monounsaturated and 50% polyunsaturated fats. K. nigricans (9%) has higher total fat/oil content than K. coccinea (3%) (Rivett et al. 1983)(Table 2). Kennedia prostrata seeds contain >22% protein (N% × 5.7), which is concentrated in the embryo and cotyledon (Hocking 1980). The embryo and cotyledons made up only 23.7% of the seed weight compared to the testa (seed coat) which made up 75% of seed weight and contained over 30% of its N and P (Hocking and Kortt 1987). 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%, Lupinus 17.5%), and

28 contains <5% of the seed’s N and P (Hocking 1980). Hence, it appears that significant gains could be made in improving total protein yield from Kennedia seed by selecting for a 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 observed in other Kennedia (over 95% of seeds were dormant at maturity in K. rubicunda), which causes problems for uniform and reliable germination 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 embryo (Smartt 1976).

Plate 27. Leaves and pods of a Kennedia eximia plant growing in a nutrient poor soil near Gnowangerup in the south of the wheatbelt of Western Australia, showing prostrate habit and curled remains of pods after shattering

Plate 28. Ruby red flowers of Kennedia eximia growing in a glasshouse

Plate 29. Vivid purple flowers of Kennedia prorepens growing in a glasshouse

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Plate 30. Large, red flowers of K. prostrata growing in the glasshouse

Plate 31. Despite possessing the prostrate habit common in the genus, K. prorepens produces flowers and pods on erect peduncles, separating them from the canopy and making harvest potentially much easier

Plate 32. The size and habit of these two collections of K. prostrata grown under identical conditions demonstrate some of the diversity present in this widely distributed species

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Plate 33. Kennedia prostrata can be tolerant of very sandy, nutrient poor soils such as the coastal dunes at the germplasm collection site in the foreground

Plate 34. Despite producing masses of herbage, these Kennedia prostrata plants did not flower until they were two years of age

Plate 35. Kennedia eximia seed

Plate 36. Kennedia prorepens seed

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Plate 37. These two samples of seed, varying greatly in size, came from two collections of the same species, Kennedia prostrata

Although no major toxicity problems have been documented with Kennedia, Rivett et al. (1983) found K. nigricans and K. coccinea seeds to contain significant concentrations of canavanine, 8.1 and 6.0 mol%, respectively. However, the presence of canavanine in some of these seeds may not prove an obstacle to their food use since the apparent toxicity of this compound is low.

Overall, Kennedia is 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 favourable 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 prioritised for further investigation (Table 3). Germplasm of K. prostrata may also have some desirable adaptation to challenging environments, but its inability to flower and set seed in its first year is considered to be a major constraint (Table 3).

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(a)

(b)

(c)

Figure 4. Australian distribution of native (a) Rhyncosia, Glycyrrhiza and Hardenbergia spp., (b) Indigofera spp., and (c) Kennedia spp. mapped against targeted agro-climatic regions; semi-arid cropping zone (E2, E3, E4) – light grey; and the arid interior (E6, G) – dark grey (see Fig. 1). Data sourced from Australian Virtual Herbarium (Council of Heads of Australian Herbaria 2008)

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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 grain crops (e.g. L. corniculatus, L. pedunculatus, L. glaber). The genus is large, but only two species are native to Australia, L. australis and L. 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 be better adapted 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 resistant (Cunningham et al. 1981).

Both Australian Lotus species can perenniate and flower in their first year, although L. cruentus often acts as an annual and produces copious seeds (Moles et al. 2003). L. australis has an erect–ascending habit growing to 60 cm in height, while L. cruentus is more prostrate to ascending (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010). Flowering can occur all year round but mainly in spring with maturity in early- to mid-summer (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010). Plants are pollinated by insects, commonly bees, although L cruentus appears to be reasonably self-compatible (Richard Bennett, unpublished data) (see also Plates 38–45).

A major limitation of Lotus species for grain production is loss of seed, due to their continuous flowering and high propensity for pod shatter at maturity. Seed shattering (dehiscence) is a major problem for seed production in domesticated Lotus species used as forage plants and seed losses vary between 5 and 88% (Garcia-Diaz and Steiner 2000). Lotus species are generally small seeded, with seeds weighing between 1.3 and 2.7 g (Table 1; Plates 46–47). Seed are often smooth, very round and brown coloured with 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 sheep (Cunningham et al. 1981; Jermyn 1985). However, significant variability in HCN content has been identified in L. australis, enabling genotypes with low HCN to be selected and bred (Real et al. 2005). Highest concentrations of HCN are in leaves and flowers, with much lower concentrations in seeds and pods which decrease as pods mature (Gebrehiwot and Beuselinck 2001). Thus, HCN is unlikely to be a major limitation for Lotus species and could be addressed through breeding, if required.

Australian Lotus species have attracted interest as potential forage legumes for low rainfall environments (Real et al. 2005), but they seem less suited as grain crops. They have quite small seeds, which are ordinary in appearance so would have limited 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 may have limited suitability as grain crops (Table 3).

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(a)

(b)

(c)

Figure 5. Distribution of Australian native (a) Lotus and Trigonella spp., (b) a selection of widely distributed Swainsona spp., and (c) native Vigna spp. mapped against targeted agro-climatic regions; semi-arid cropping zone (E2, E3, E4) – light grey; and the arid interior (E6, G) – dark grey (see Fig. 1). Data sourced from Australian Virtual Herbarium (Council of Heads of Australian Herbaria 2008)

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Plate 38. Despite their size, these tiny Lotus australis plants growing on a nutrient impoverished coastal sand dune are still producing flowers

Plate 39. The thick, almost succulent, leaves of this Lotus australis plant may help it cope with salt exposure from the nearby ocean and periodic drought conditions on freely draining soils

Plate 40. The copious pods and almost complete lack of functioning leaves are a demonstration of the reproductive ability of these Lotus australis plants

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Plate 41. A Lotus australis plant displaying an ideal stature for harvest using a conventional header

Plate 42. The many orange brown pods in the foreground are produced by Lotus australis plants growing on a sand dune just metres from the ocean

Plate 43. An alternative form of Lotus australis (grey/green plants in foreground) under investigation for its potential to provide fodder for livestock under very dry conditions

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Plate 44. Flower of Lotus cruentus in the glasshouse

Plate 45. Lotus cruentus can have excellent fecundity and a short generation time

Plate 46. Lotus australis seed (accession PD01LA)

Plate 47. L. australis seed (accession SA37841)

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Rhynchosia

Rhynchosia includes more than 200, mostly tropical, species with six native to Australia. Several species in the genus are commonly called rosary bean because of their attractive red, blue, black, mottled or bicoloured seeds. The pantropical species R. 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 species (Harding et al. 1989).

Most Australian Rhynchosia are restricted to sub-tropical 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 soils (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010; Harding et al. 1989). However, it has been collected from sands and sandy loams (Harding et al. 1989). 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 an abundance of seed (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010; Harding et al. 1989). Days to flowering vary from 43 to 142 (Harding et al. 1989). Pod indehiscence was observed in most accessions of Rhynchosia, but some accessions retain seeds longer than others (Harding et al. 1989).

Seeds of R. minima are reasonably large (9–12 mg), kidney-shaped and greyish, brown or black and often mottled. However, seed size ranges from 8.8 mg to 20.4 mg (Harding et al. 1989). There is no information on the concentrations of protein or oils in R. minima seed, but they contain some chemicals of pharmaceutical interest including prodelphinidin (antibiotic), gallic and protocatechuic acid (antiasthmatic, antioxidant) (GRIN (Germplam Resources Information Network); Morris 1997).

R. minima warrants further appraisal as a grain crop (Table 3; Plates 48–53). 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, pod indehiscence and seed size providing potential to identify and select desirable genotypes to improve grain production. Its distribution suggests it is tolerant of water-limited environments, but its preference to heavy clay soils may restrict its application in some regions.

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Plate 48. These juvenile Rhynchosia minima plants are already displaying their strong twining habit

Plate 49. Pods of Rhynchosia minima, each containing a maximum of two large seeds

Plate 50. Pods and flowers of these Rhynchosia minima plants grown without climbing frames are produced on erect peduncles

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Plate 51. A massive, symbiotic nitrogen- fixing root nodule taken from the root of a Rhynchosia minima plant sectioned to show the red, actively n-fixing tissue. Herbaceous legumes that produce nodules of this size are rare

Plate 52. This four day old seedling of Rhynchosia minima is demonstrating excellent vigour conferred in part by the large seed, with the root growing more than 2 cm each day

Plate 53. Rhynchosia minima seed

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Swainsona

Swainsona includes 85 species of which 84 are endemic to Australia. The best known of these is S. formosa (Sturt's Desert Pea) which is grown as a 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 more widely distributed species include S. formosa, S. canescens, S. colutoides and S. swainsonoides (Fig. 5b). Many species also exhibit characteristics of plants adapted to dry environments such as hair covered 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 sub-shrubs that may range from prostrate to semi-erect (Florabase - The Western Australian Flora 1998). At least a few species are winter-growing that flower and set seed in spring (e.g. S. canescens). Most species seem to be predominantly open pollinated by insects or birds (Florabase - The Western Australian Flora 1998), but some have a degree of self-compatability. A few species are known to exhibit exceptional fecundity, for example, well grown plants of S. canescens are capable of setting approx 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 indehiscence species (e.g. S. canescens, S. colutoides, S. pyrophila, S. fraseri). Yet, pods may still senesce with seeds enclosed. Seeds of Swainsona are small (usually <3 mg) and kidney-shaped, with surprisingly little variation in size between the species. Seeds usually have hard seed coats, which induce dormancy.

At least one Swainsona species, S. galegafolia (Darling pea), was used by Aborigines, and was eaten green with a similar taste to the common garden pea (Brand and Cherikoff 1985). Analysis of green seed (69% moisture content) found it to contain 31% protein, 33% carbohydrate, 26% fibre and 6% fat (on a dry matter basis) (Brand and Cherikoff 1985). However, they also recorded that 64% of the seed was edible (by mass), but 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 swainsonine (Jermyn 1985); while this is poisonous to livestock, its effect on humans is unknown. Aplin and Cannon (1970) reported that the concentration of alkaloids in general (not only swainsonine) in the vegetative material of other Swainsona species was high in S. rostellata, moderate in S. campestris, S. canescens, S. incei, S. stipularis and low in S. cyclocarpa, S. flavicarinata, S. occidentalis. Species reputedly or proven to be poisonous when grazed by livestock include S. galegifolia, S. sejuncta, S. greyana, S. lessertiifolia, S. luteola, S. microphylla, S. oroboides and S. procumbens (Gardiner et al. 1969). Swainsonine is found in the seeds of some Swainsona: S. galegifolia and S. sejuncta contain 2900 and 1700 mg/kg of swainsonine, respectively (Martyn et al. 2003). There is little 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 stems or leaves of S. galegifolia (up to 7500 mg/kg) and S. sejuncta (up to 5200 mg/kg) is approximately 2.5 times the concentration in the seeds (Martyn et al. 2003). Thus species, such as S. formosa which have low concentrations of swainsonine in leaves (70 mg/kg) and flowers (210–490 mg/kg) may have very low concentrations in seeds (Martyn et al. 2003).

Despite the lack of information on many Swainsona species, a number have characteristics that suggest they are worthy of further investigation for their grain production potential (Table 3; Plates 54–66). 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, S. purpurea and S. kingii) for which little information was available could also possess desirable attributes.

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Plate 54. Several Swainsona canescens in the wheatbelt of Western Australia near Newdegate

Plate 55. Swainsona canescens plants grown under irrigation produced masses of white fluffy pods that may be easily harvested with a header

Plate 56. The large, elegantly coloured inflorescences of Swainsona canescens led to interest in its development as a horticultural plant

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Plate 57. Swainsona canescens plants growing in their natural habitat near Mount Magnet, Western Australia, and producing erect inflorescences covered in white woolly pods

Plate 58. The large purple flowers of Swainsona colutoides with red immature pods in the background

Plate 59. A Swainsona colutoides plant with red immature and brown mature pods held well on the plant; this would enable easy harvest with conventional machinery

Plate 60. These Swainsona formosa (Sturt’s Desert Pea) plants produced their iconic flowers well under irrigated conditions but pods and seeds were rare

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Plate 61. These small Swainsona kingii plants can produce many pods and seeds despite their growth being limited by environmental conditions

Plate 62. A Swainsona purpurea plant showing its nitrogen-fixing root nodules

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Plate 63. Swainsona purpurea does not produce seed in the absence of pollinators and requires bees or other insects to ‘trip’ flowers, breaking a membrane on the stigma to allow pollination to occur

Plate 64. Swainsona canescens seed

Plate 65. Swainsona kingii seed

Plate 66. Swainsona purpurea seed

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Trigonella

Of the 80 species in the genus Trigonella, T. suavissima (sweet fenugreek) is the only native of Australia (Legume Web - ILDIS World Database of Legumes vers. 10 2008). This species has been investigated for its potential as a forage plant and possesses many suitable agronomic traits (Halloran and Pennell 1981). Several exotic Trigonella are important for culinary, nutritional or medical reasons (Acharya et al. 2006). The most widely used is fenugreek (T. 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 manured (Hymowitz 1990; McCormick et al. 2009).

T. suavissima is a winter-growing annual or ephemeral, flowering between autumn and spring (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010). It occurs throughout inland arid environments in central Australia, where it is typically found on heavy clay soils of river banks, floodplains and depressions (Cunningham et al. 1981)(Fig. 5a). It is rarely found on sandy soils (Cunningham et al. 1981). 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 lifecycle may allow it to avoid severe water stress rather than tolerate water deficit. T. suavissima has tolerates salinity well compared to other native and exotic legumes, with growth of 106% of control at NaCl concentrations of 40 mM (Rogers et al. 2005). T. sauvissima has a desirable growth habit being decumbent to ascending and reaching 50 cm in height (PlantNET - The Plant Information Network System of Botanic Gardens Trust 2010). Little agronomic data is available on the seed production potential of T. sauvissima, yet it is reputed to have a high level of fecundity (Cunningham et al. 1981). Collected accessions of the species flowered between 111 and 118 days after sowing. It is commonly pollinated by insects, but its self- compatibility is unknown (Dear et al. 2003). A favourable attribute of T. sauvissima is that it can be indehiscent or tardily dehiscent, yet fruits are often shed from plants at maturity (Florabase - The Western Australian Flora 1998; Dear et al. 2003).

The seeds of one tested accession of T. suavissima are small (1 mg, Table 1; Plate 67); substantially smaller than its grain legume relative fenugreek (9–22 mg) (McCormick et al. 2009). The small seeds of T. suavissima may limit its yield potential as a grain crop. However, it may have use as a multipurpose pasture and crop species (as for fenugreek), as it is regarded as a valuable and nutritious fodder source where it grows naturally (Halloran and Pennell 1981). There are no published studies of seed chemical composition or presence of bioactive compounds in T. suavissima. Fenugreek contains a number of bioactive chemicals and has beneficial medicinal and nutritional qualities (Bordia et al. 1997). The presence of these qualities in T. suavissima is worth exploring. Overall, Trigonella suavissima deserves 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 may potentially contain chemicals with pharmaceutical applications. The major limitation appears to be small seed size but exploration of variability in this trait may reveal germplasm with greater seed size. More information is required on the seed chemistry and seed yield potential of this species.

Plate 67. Small seed ofTrigonella suavissima

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Vigna

The Vigna genus contains a number of species that are widely grown as grain legumes throughout the world (e.g. mung bean (V. radiata), azuki bean (V. angularis) and cowpea (V. unguiculata)) and some secondary grain legumes (e.g. V. acontifolia (moth bean), V. lanceolata (pencil yam), V. mungo (urad bean, black gram), V. subterranea (Bambara groundnut), V. umbellata (rice bean), and V. vexillata (zombi pea)) (Hymowitz 1990). Five species of Vigna are indigenous to Australia and one is endemic (V. lanceolata) (Lawn and Watkinson 2002). V. radiata ssp. sublobata is the putative progenitor to the cultivated mungbean (V. radiata) and is a native of Australia (Legume Web - ILDIS World Database of Legumes vers. 10 2008).

Australian Vigna species are predominately tropical or found mainly 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 which exhibit significant variation in important agronomic traits (e.g. seed yield, days to flowering, frost tolerance) (Lawn and Holland 2003). 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 and twining habit, and lower seed yield and harvest index (Grant et al. 2003; Lawn and Rebetzke 2006; Rebetzke and Lawn 2006). Overall, Australian Vigna are of secondary interest because of their predominantly tropical distribution but others have investigated their agronomic potential (Grant et al. 2003; Lawn and Rebetzke 2006; Rebetzke and Lawn 2006).

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Conclusion

Australia has a diverse flora of herbaceous legumes and their agricultural potential, especially their potential to produce grain products, has been little assessed. For many species, data is sparse and must be viewed carefully since it is likely that high variability within individual species has not been encompassed in past studies. However, many species possess characteristics useful in marginal grain growing environments due to their adaptation to arid and semi-arid climates and, sometimes, infertile soils. A major challenge for some species (e.g. Glycine, Kennedia, Rhychosia) is their twining growth habit. However, this was the case in many progenitors of modern legume grain crops. Similarly, substantial increases in seed size and removal of seed dormancy mechanisms have been achieved through plant breeding, so while many species have small seeds, there may be potential to increase their size. Little information exists on the chemical constituents of many native Australian legumes but some may find a market because they possess attractive seeds (especially for small seeded species e.g. Glycine, Glycyrrhiza) or because they possess bioactive compounds with potential for natural medicinal uses (e.g. Glycine, Cullen, Trigonella, Indigofera). While germplasm of native legumes is currently stored in Australian Genetic Resource Centres (e.g. see Hughes et al. 2008), it is unlikely that these collections fully represent the diversity present in natural populations and any serious attempt at domestication of most native legumes would probably need to commence with a comprehensive collection of wild germplasm (Bennett et al. in press).

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Table 3. Prioritisation of species for further investigation as grain legume crops. Species with little information are not included (? – indicates where information is lacking)

Distrib. Life C Seed Flowers Pollinatio Pod/seed Bioactivity Species A B Habit D Other notable qualities/information cycle size 1st year n retention considerations Highest priority species for further investigation Cullen tenax **** P Sp, C Small Y Selfing Low Furanocoumarins Aerial seed Crotalaria cunninghamii **** P E Large ? ? ? - Pods on branch ends Glycine canescens **** P T Mod. Y Selfing Low Phytoestrogens Glycyrrhiza acanthocarpa **** P E, C Mod ? ? High - Salt tolerance, Pods easy to thresh Mod– Kennedia prorepens **** P T Y Open Low - Large Mod- Rhynchosia minima *** P T – SE ? Selfing Variable - Large Swainsona canescens **** A/B SE, Sp Small Y Open Mod-High Swainsonine Mod, delayed Swainsona colutoides *** A/B E Small Y Selfing Swainsonine Pod retention is good, inflated pods indehiscence Trigonella suavissima **** A E – SE Small Y ? ? - Good waterlogging & potential salt tolerance Moderate priority species with a number of suitable attributes Cullen australasicum **** P E Mod Y Open Variable Furanocoumarins

50 Small– Cullen cinereum *** A/B E, Sp Y Selfing Variable Furanocoumarins mod

Cullen graveolens *** A/B E – SE ? Y ? ? Furanocoumarins Open, Cullen pallidum *** P Sp ? Y some Variable Furanocoumarins selfing Glycine latifolia ** P T Mod Y ? Phytoestrogens Systematically collected/ selected as forage Small– Glycine tabacina ** P T Y ? Low Phytoestrogens mod. Kennedia coccinea * P P, T Large Y ? ? - Indigofera australis *** P E, Sp Mod ? ? Indospicine, HCN Swainsona formosa **** A SE, Sp ? Y ? ? Swainsonine Swainsona *** P Sp Small Y Open Low Swainsonine swainsonioides Species with some valuable attributes but some limitations Kennedia prostrata ** P P, T Large N Open - Kennedia nigricans * P P, T Large ? Open - Lotus cruentus **** A Sp, C Small Y Selfing Very low HCN A Match between species distribution and targeted agro-climates: **** highly favourable, *** favourable, ** moderate, * poor B P: perennial, A: annual, B: biennial C E: erect, SE: semi-erect, P: prostrate, T: twining/trailing, Sp: spreading, C: clumping/crown forming D Large: 10–20 mg, mod: 5–10 mg, Small: <5 mg

Section 2. Glasshouse trial

Summary 17 accessions of native Australian herbaceous legumes were assessed in the glasshouse over autumn/winter for 20 growth traits and seed nutritive value. Eight traits related to domestication and harvestability (days to flowering, days to podding, harvest index, seed yield/plant, duration of flowering, vegetative dry weight, 1000 seed mass, and height at harvest) showed significant differences across species. Three commercially available grain legume cultivars chickpea cv Rupali, field pea cv Kaspa, and narrow leaf lupin cv Mandelup were included in the study. The major differences between the native species and the commercial cultivars were as follows:

• Native species generally took longer to begin flowering and the duration of flowering was much longer, with eight species flowering continuously once flowering commenced.

• Harvest index, seed weight/plant and seed size were generally much lower for the native species.

• Days to podding was generally higher for the native species, with the exception of Rhynchosia minima, Swainsona colutoides and Trigonella sauvissima which were similar to field pea, and Lotus cruentus and S. kingii which were similar to lupin.

• Above-ground vegetative dry weight of native species was generally higher than for the commercial cultivars, with the exception of L. cruentus and S. kingii which produced very little vegetative dry mass.

• Most native species were low growing with a similar height to chickpea at harvest. However, Glycine canescens was much taller and was of a similar height to the field pea, and G. sp. and Kennedia prorepens were of intermediate height.

• Nutritive value of seed was generally similar to that of the commercial cultivars.

• Fat content of native legume seed ranged between 1.9% and 5.7% with exception of Cullen australasicum and C. tenax which had elevated fat content at 10.7% and 10.4%, respectively.

• Protein content was highest for T. sauvissima (33.6%) which was very similar to lupin (33.2%). All other native species with the exception of R. minima (protein content of 18.8%) had protein contents within the range of the three commercial cultivars (20.3% - 33.2%).

• Dietary fibre was also within the range for three commercial cultivars (13.4% - 42.6%), with most native species within the range 18.8% - 31.7%. The highest dietary fibre content was found in T. sauvissima (38.6%) which is lower than the highest commercial cultivar lupin cv Mandelup (42.6%).

These results indicate the native species have some of the characteristics required for domestication and have similar nutritional values to existing crops.

Introduction In Section 1 of this report, a review of the literature was used to evaluate the potential of herbaceous species from 14 genera of Australian legumes from arid and semi-arid climatic regions for their suitability as grain crops. Species were classified into one of three priority ratings for further

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investigation based on the presence of characteristics suited to broad acre farming as grain legumes or bioactive compounds with potential as natural medicines (Table 3). It was acknowledged that the prioritisation process was hindered by a lack of both qualitative and quantitative data on seed qualities and plant growth characteristics. A glasshouse experiment was therefore conducted to collect this data for some of the higher priority taxa.

Materials and Methods Priority species for testing were selected based on the results of unpublished evaluations of their potential as pasture legumes by Steven Hughes, SARDI Genetic Resource Centre, Waite Research Precinct, Plant Research Centre, 2b Hartley Grove, Urrbrae SA 5064, and of published information on characteristics related to harvestability as summarised in Table 3. The 17 native legume species included in the study are listed in Table 4. Where seed of high priority species were unavailable, substitutions were made with lower priority species. Seed was obtained from the following sources: SARDI Genetic Resource Centre, Waite Research Precinct, Plant Research Centre, 2b Hartley Grove, Urrbrae SA 5064 Australia; Australian Tropical Crop and Forage Resource Centre, Department of Primary Industries and Fisheries, Biloela Research Station, State Farm Rd, Biloela, Qld. 4715 Australia; Kimseed International, 4/5 Collingwood St., Osbourne Park, WA 6017, Australia; Nindethana Seed Service Pty. Ltd., PO Box 2121, Albany, WA, 6331, Australia, and collections made throughout the arid and semi-arid agricultural regions of Western Australia as part of various projects run by the CRC for Plant-based Management of Dryland Salinity.

Seeds of all species were scarified, imbibed on dampened filter paper and sown the following day in mid-May 2008. Six seeds of each species were sown per pot in ten replicate 300 mm black standard pots with drainage holes filled with UWA potting mix. UWA potting mix is prepared from the following bulk ingredients; 2.5 m3 fine composted pine bark, 1 m3 coco peat royered, 1.5 m3 brown river sand, 5 kg superphosphate, 10 kg extra fine limestone, 1.5 kg potassium sulphate, 1 kg macro mineral trace elements, 5 kg ammonium nitrate (Agran 34.0), 10 kg dolomite (CalMag) and 2.5 kg ferrous sulphate heptahydrate. Pots were placed in a glasshouse at ambient temperature and watered every Monday, Wednesday and Friday (less frequently if pots were still moist) and fertilised every 2 weeks with approximately 100 ml of a solution of Phostrogen fertiliser (Phostrogen Ltd., Deeside, Clwyd, CH5 2NS distributed in Australia by Debco Pty Ltd, 12 McKirdy’s Rd., Tyabb, Victoria, 3913) made up at the rate of one scoop (10 ml) dissolved in 10 litres of water. Thus it was expected that plant growth would not be limited by either nutrients or water. Seedlings were thinned to 3 per pot on 20th June 2008 (7 weeks after sowing). Watering of commercial cultivars ceased 26 weeks after sowing (10th November 2008) and of native species 31 weeks after sowing (15th December 2008). Native species were harvested later because of a later start to flowering and pod set.

Plants were harvested within 3 weeks of cessation of watering. Roots were washed carefully and plants separated into pods and seeds, shoots, and roots. Shoots and roots were dried in an oven at 70oC for 72 hr and dry weights recorded. Seeds were air dried, weighed and stored at room temperature prior to analysis. Data were collected for the following traits: days to emergence; days to flowering; number of nodes from soil level to first flower on main stem; number of flowers per inflorescence; height at first flowering; length of main stem at first flowering; number of primary and secondary branches at first flower; extent of branching at first flowering; growth habit; days to first pod; mean number of pods per inflorescence; first pod node; number of days to end of flowering; duration of podding; days from sowing to maturity; shattering; height at maturity; branching at maturity; dry weight shoots; dry weight roots; dry weight total; number of pods at harvest; number of seeds at harvest; seed weight. Harvest index (%) was calculated as = 100 X (Seed dry mass (g)/Plant dry mass (g)). N.B. Only vegetative data was available for species which did not flower. Only characteristics related to harvestability showing significant differences across species are reported. Lupin plants became diseased during the experiment and died early, thus stored seed was used for analysis.

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Nutrient analysis of seed was carried out by George Weston Food Technologies for fat, protein and dietary fibre. The analytical methods: Moisture - AOAC 930.15, 925.10, 950.46, 925.45; Protein (Combustion) - AOAC 990.03, 992.15, 992.23, 993.17, 997.09; Fat Soxhlet Extraction - AOAC 920.39; TDF - AOAC 985.29 (Phosphate Buffer) were taken from the 17th edition of Official Methods of Analysis of AOAC International (Association of Official Analytical Chemists International).

Growth data were analysed by general analysis of variance (ANOVA) in Genstat version 10 (Lawes Agricultural Trust, Rothamsted Experimental Station, UK, 2007). The l.s.d. values at P=0.05 are presented.

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Table 4. The native Australian herbaceous legumes used in the study. The commercially available cultivars; Cicer arietnum cv Rupali (chickpea), Pisum sativum cv Kaspa (field pea), and Lupinus angustifolius cv Mandelup (narrow leaf lupin) were also included. Species, authority * Accession number Growth habit Life cycle Priority ranking

Cullen australasicum (Schltdl.) J.W.Grimes ID42 SA44380 erect perennial 3 Cullen cinereum (Lindl.) J.W.Grimes ID51 AusTrCF 320112 erect annual/biennial 2 Cullen graveolens (Domin.) J.W.Grimes ID76 AusTrCF 320184 erect annual/biennial 1 Cullen tenax (Lindl.) J.W.Grimes ID97 AusTrCF 320110 erect perennial 1 Glycine canescens F.J.Herm NIND001 climber perennial 1 Glycine sp. NIND004 climber n.a. n.a. Glycyrrhiza acanthocarpa (Lindl.) J.M.Black C2N01GA erect perennial 1 Kennedia coccinea Vent. NS-26828 prostrate perennial 2 Kennedia prorepens F.Muell NS-30323 prostrate perennial 1

54 Lotus cruentus Court NF003 prostrate annual 2 Rhynchosia minima (L.) DC. NF013 prostrate perennial 1 Swainsona canescens (Benth.) F.Muell. KIMS003 erect annual/biennial 1 Swainsona colutoides F.Muell. NIND006 erect annual/biennial 1 Swainsona kingii F.Muell. NF002 prostrate annual/perennial n.a. Swainsona purpurea (A.T.Lee) Joy Thomps. KIMS004 semi-erect annual/perennial 3 Swainsona swansonioides (Benth.) J.M.Black KIMS005 semi-erect perennial 2 Trigonella sauvissima Lindl. n.a. erect annual 1 * Authority obtained from Florabase, http://florabase.calm.wa.gov.au/ , accessed May 2010

n.a. = not available

Table 5. Eight plant growth traits measured in the glasshouse experiment. Predicted means and LSD (p=0.5) from ANOVA. Values followed by the same postscript are not significantly different.

Duration of Days to Days to Seed yield Vegetative dry 1000 seed Height at Species Harvest index flowering flower podding (g/plant) weight (g) mass (g)xx harvest (mm) (days) Cullen australasicum 132 gh 164 h 0.093 ab 4.25 bc 68 d 42.7 f 8.7 c 330 ab Cullen cinereum 115 ef 177 ij 0.304 d 6.18 cd dnf 13.5 de 5.2 def 363 ab Cullen graveolens 122 fg 161 h 0.106 b 2.45 ab dnf 15.8 cd 5.7 de 301 ab Cullen tenax 116 ef 171 hi 0.295 cd 8.32 de dnf 17.7 cd 5.2 def 491 bc Glycine canescens 157 h 182 ij 0.350 d 8.05 de dnf 15.5 h 16.9 a 1366 e Glycine species 104 d 172 hi 0.543 ef 14.35 f 79 e 12.3 a 11.2 b 702 cd Glycyrrhiza acanthocarpa 202 i 211 k na na dnf 15.6 cd na 444 ab Kennedia coccinia dns na na na na na na na

55 Kennedia prorepens 167 h 187 j 0.001 a 0.05 a dnf 41.0 fg 6.3 cd 782 d Lotus cruentus 81 bc 88 bc 0.650 g 10.20 e 100 g 5.6 abc 1.5 g 340 ab Rhynchosia minima 107 de 120 f 0.065 ab 2.59 ab 84 e 37.7 h 13.3 b 497 bc Swainsona canescens 130 g 144 g 0.016 ab 0.50 a dnf 29.4 2.7 g 425 ab Swainsona colutoides 110 de 112 ef 0.208 c 6.02 cd 83 e 22.7 fg 3.1 fg 438 ab Swainsona kingii 92 c 99 cd 0.474 e 6.53 cd 91 f 7.3 bcd 2.7 g 245 a Swainsona purpurea 111 de 138 g 0.004 a 0.13 a 69 d 31.5 h 3.2 efg 463 ab Swainsona swansonioides 111 de 161 h 0.001 a 0.04 a dnf 40.6 gh 5.5 def 528 bc Trigonella sauvissima 74 b 104 de 0.345 d 4.30 bc 101 g 10.6 bcd 1.2 g 444 ab Chickpea cv Rupali 60 a 65 a 0.601 fg 13.68 f 44 c 9.4 ab 188.7 xx 457 ab Field pea cv Kaspa 109 de 115 ef 0.495 e 29.83 g 28 b 31.2 abc 258.9 xx 1077 e Lupin cv Mandelup 74 b 81 b deaf dead 14 a dead dead dead LSD (0.05) 11.1 11.3 0.09 2.811 6.7 6.52 2.44xx 232.5 na - not available; dns – did not flower; dnf – did not finish flowering xx – significance analysis conducted without chickpea and field pea data

Results Eight traits related to domestication and harvestability (days to flowering, days to podding, duration of flowering, vegetative dry weight, height at harvest, harvest index, seed yield/plant, and 1000 seed mass, showed significant differences across study species.

Flowering was generally initiated much later in the native species than in the commercial cultivars, at 74 to 202 days post planting compared with 60 to 109 days (Table 5). However, five species were within the range for the commercial cultivars; Glycine sp., Lotus cruentus, Rhynchosia minima, Swainsona kingiii and Trigonella sauvissima. Days to podding ranged between 88 days for L. cruentus to 211 days for Glycyrrhiza acanthocarpa. These values were generally higher than for the commercial cultivars with the exception of R. minima, S. colutoides and T. sauvissima which were similar to field pea, and L. cruentus and S. kingii which were similar to lupin (Table 5). Pod shattering characteristics were scored for 10 of the native species and two of the commercial cultivars. Six of the native species had mild or severe shattering of pods (Table 6).

Table 6. Shattering characteristics of 10 native herbaceous legumes and two commercial cultivars. Intact pod drop was not recorded as shattering. Full seed retention Mild shattering Severe shattering Cullen australasicum Glycine canescens Trigonella sauvissima Cullen cinereum Glycine species Cullen graveolens Lotus cruentus Cullen tenax Swainsona canescens Chickpea cv Rupali Swainsona purpurea Field pea cv Kaspa

Cullen australasicum, G. sp., Kennedia coccinia, L. cruentus, R. minima, S. colutoides, S. kingii, S. purpurea and T. sauvissima had a distinct flowering period and had completed pod set by harvest. Only two of these species, L. cruentus and T. sauvissima, are described in the literature as annuals, while the remainder are annual/biennial or annual/perennial (no information is available for currently unidentified G. sp.) (Table 4). A number of biennial or perennial taxa; C. cinereum, C. graveolens, C. tenax, G. canescens, Gl. acanthocarpa, K. prorepens, S. canescens and S. swansonioides were still flowering when watering ceased (Table 5). In comparison, all of the commercial cultivars had distinct flowering and pod set periods which were completed before watering of native species ceased (Table 5).

Above-ground vegetative dry weight of native species was generally higher than for the commercial cultivars, with the exception of L. cruentus and S. kingii which produced very little vegetative dry mass (Table 5). Most native species were low growing (range 245 to 528 mm) with a similar height to chickpea (457 mm) at harvest (Table 5). However, G. canescens was much taller (1366 mm) and was higher than the tallest commercial cultivar, field pea at 1077 mm. G. sp. and K. prorepens were of intermediate height, 702 mm and 782 mm, respectively (Table 5).

Seed yield per plant was generally low for native species with only two species, Glycine sp. (14.4 g/plant) and L. cruentus (10.2 g/plant) producing yields close to that of chickpea (13.68 g/plant). The seed yield of chickpea was less than half of that of field pea (29.8 g/plant) (Table 2). Seed size, determined by measuring 1000 seed mass, was very low for all native species with the highest value 16.9 g (G. canescens) less than 10% of the value for the smallest commercial cultivar seed (chickpea, 188.7 g.) Harvest index (seed mass as a percentage of total plant dry mass) was very low for S. canescens, S. purpurea and S. swansonioides (0.016, 0.004 and 0.001 %) and generally lower for the

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native species than commercial cultivars with the exception of G. sp., L. cruentus and S. kingii (Table 5).

Nutritive value of the seed of the native species was generally similar to that of the commercial cultivars. Fat content of native seeds ranged between 1.9% and 5.7% with the exception of C. australasicum and C. tenax which had an elevated fat content at 10.7% and 10.4%, respectively (Table 7). Protein content was highest for T. sauvissima (33.6%) which was very similar to lupin (33.2%). All other species with the exception of R. minima (protein content of 18.8%) had protein contents within the range for three commercial cultivars (20.3% - 33.2%) (Table 7). Dietary fibre was also within the range for the controls (13.4% - 42.6%), with most species within the range 18.8% - 31.7% (Table 7). The highest dietary fibre content was found in T. sauvissima (38.6%) which was, however, lower than the highest commercial cultivar, lupin cv Mandelup (42.6%) (Table 7).

Table 7. Nutrient analyses of grain of native Australian herbaceous legumes and commercially available cultivars. Species Fat Protein Dietary Fibre Moisture (%) (%) (%) (g/100g) Cullen australasicum 3.4 29.2 25.8 9.8 Cullen cinereum 10.7 32.8 23.8 9.3 Cullen graveolens 5.7 30.4 n.a. 8.9 Cullen tenax 10.3 29.2 25.4 8.9 Glycine canescens 5.6 30.7 28.6 9.8 Glycine sp. 4.7 29.1 31.7 9.5 Glycyrrhiza acanthocarpa n.a. 24.8 n.a. 4.9 Kennedia coccinea n.a. n.a. n.a. n.a. Kennedia prorepens n.a. n.a. n.a. n.a. Lotus cruentus 5.3 28.8 38.6 10 Rhynchosia minima n.a. 18.8 n.a. 10.4 Swainsona canescens 3.6 24.5 20.7 9 Swainsona colutoides 1.9 24.6 21.4 10.5 Swainsona kingii 2.3 31.3 21.5 8.8 Swainsona purpurea n.a. n.a. n.a. n.a. Swainsona swansonioides 3.3 30.1 18.1 7.5 Trigonella sauvissima 4.8 33.6 18.4 9.8

Chickpea cv Rupali 4 20.3 22.8 11.4 Field pea cv Kaspa 1.1 23.3 13.4 11.5 Lupin cv Mandelup 4.6 33.2 42.8 10 n.a. = not available (insufficient seed for analysis)

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Discussion The results of the glasshouse study indicate that all the native herbaceous species that were evaluated had some of the characteristics required for domestication and had similar nutritional values to existing commercial grain legume cultivars.

The results of the experiment suggest that the major limits to domestication of native legumes may be lack of determinate flowering, low harvest index and low seed size. All of these characters have been improved easily in the domestication of other exotic legumes as grain crops (Fuller 2007; Weeden 2007). Furthermore, the end-use of a native grain may mean that some standard characteristics of exotic grain legumes may not be required. For instance, a native legume may attract a price premium due to novelty value and thus yield may not need to match that of exotic grain legumes. Alternatively, adaptation to low fertility soils and drier climates may provide a niche for natives where they are not in competition in terms of yield with current commercial grain legume species.

While no one native species is suggested by the results of the glasshouse trial as an obvious focus for further research, efforts should be made to identify the accession labelled as Glycine species, as it showed a number of promising characteristics, notably high seed yield and harvest index, and high seed protein and dietary fibre contents.

A major limitation of the glasshouse study was the inclusion of only one accession of each species. When a large number of accessions of C. australasicum were assessed for pasture potential in the wheatbelt of Western Australia, large variation was found between accessions in many characters (Bennett et al. 2006). Further work needs to be directed towards assessing a larger number of accessions to determine the variability of important characteristics within each species. A useful first step would be to follow the methodology of Bennett et al. (in press) who used an analysis of soils and climate niche models of herbarium and germplasm records of the 16 perennial, herbaceous Australian Cullen species. The study identified which species were suited to the target area, in this case the drier regions of the Western Australian wheatbelt, but also found that the existing germplasm collections were unlikely to well encompass the variation present in wild populations. The degree of variability in wild populations may determine the ease with which simple selection may be used to improve characters.

A further area meriting attention is the chemical analysis of seed. This needs to be undertaken to determine if native species contain toxic compounds or anti-nutritional factors that may inhibit their use, but could also be used to identify species with compounds with medicinal value.

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Plate 68. Plants in the glasshouse experiment on 26 August 2008 a) Cullen australasicum, b) C. cinereum, c) C. graveolens, d) Glycine sp., e) field peas, f) lupins, g) Kennedia coccinea, h) Swainsona canescens, i) Lotus cruentus, j) Swainsona kingii, k) S. purpurpea, l) S. swainsonioides and m) Trigonella suavissima

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Plate 69. Surviving plants in the glasshouse experiment on 31 October 2008 a) Cullen tenax, b) Glycine sp., c) Kennedia coccinea, d) K. prorepens, e) Lotus cruentus, f) Rhynchosia minima, g) Swainsona swainsonioides and h) S. kingii

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Implications

The primary implication of this report is that native herbaceous legumes do have potential to be developed as grain legume crops for the drier areas of the southern grain belt but there is not a standout species that could be recommended for immediate domestication. Further research is required and should use the priority species identified in this project as a guide.

There is a scarcity of published information on these species, especially those in the genera Swainsona, Glycyrrhiza and Crotalaria. Moreover, there is a high likelihood of substantial variability within all the native species. Thus, any future research should commence with collection of additional germplasm and the evaluation of this germplasm under glasshouse and field conditions.

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Recommendations

As considerable further research and development is required before release of a commercial native grain legume cultivar, our recommendations are targeted primarily at the Australian research community (funding bodies and researchers). Recommendations are as follows:

1) Further prioritisation of the nine priority species identified in this report should be undertaken with a clear idea of the niche for which they are aimed, from both an agricultural systems and a grain marketing perspective. As such, both agronomic trials and consultation with the food and/or nutraceutical industries are essential.

2) Further prioritisation will be most effective if informed by collection and assessment of a reasonably wide range of germplasm from all prioritised species. It should also be kept in mind that we were not able to reasonably assess all species in each genus due to lack of information and there may be promising species we have not identified as a priority.

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Appendix 1

WORKSHOP AGENDA Domestication of native Australian legumes as grain crops for drying environments Tuesday June 29th UWA, Agriculture lecture theatre

BACKGROUND

Australia has a wealth of native legumes which are untapped for their agricultural potential. These species may have special application in the face of climate change and peak phosphorus as they generally are found in regions with poor soils and low and variable rainfall. The world is relying on progressively fewer crops for food security. Small advances in traits such as drought tolerance generally come at a large cost for these species. It is possible that legumes native to Australia could provide new crops with desirable traits with a minimal amount of selection/breeding or, at least, allow exploration of traits that allow adaptation to these challenging conditions. Such species may be perennials, annuals or – indeed – ephemerals. In an Australian context, native grain crops avoid the weed risk issues that come with the introduction of a new overseas species.

PROGRAM

8.30 – 9.00 Introduction to day – general context (Megan Ryan, UWA)

9.00 - 10.15 The domestication of Near Eastern legumes/crops (Shahal Abbo, The Hebrew University of Jerusalem) (1 hour + 15 min questions)

10.15 -10.45 Morning tea

10.45 – 12.25 What do we know about the potential of Australian native herbaceous legumes?

- Review of the literature and glasshouse experiment (Lindsay Bell, CSIRO Sustainable Ecosystems Toowoomba) (40 min) - Field screening trials in WA (Richard Snowball, DAFWA, WA) (30 min) - Field screening trials in SA (Megan Ryan) (15 min) - Question time (15 min) 12.25 – 1.30 Lunch (provide own)

1.30 – 2.15 How diverse and complete are our germplasm collections? Cullen as a case study (Richard Bennett, CSIRO Sustainable Ecosystems, WA. (35 + 10 min questions)

2.15 - 2.40 Adaptations of natives to low phosphorus and drought (Megan Ryan, UWA) (20 min + 5 min questions)

2.40 – 3.10 Afternoon tea

3.10 – 4.00 Commercial prospects (Dai Suter – George Weston Foods) (40 + 10 min questions)

4.00 – 4.30 Lessons learnt from domestication of lupin in WA (Jon Clements, CLIMA, UWA) (25 min questions)

4.30 – 5.00 Conclusions from day and general discussion (Willie Erskine)

5.00 CLOSE

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Appendix 2 WORKSHOP SUMMARY AND RECOMMENDATIONS

WORKSHOP: Domestication of native Australian legumes as grain crops for drying environments - Workshop summary and recommendations

Tuesday June 29th, 2010 Agriculture lecture theatre, The University of Western Australia Attendees – approximately 40 Copies of PowerPoint presentations can be obtained from Megan Ryan [email protected]

Contributors to this overview Megan Ryan, Shahal Abbo, Richard Bennett, Lindsay Bell, Jens Berger, Jon Clements, William Erskine, Richard Snowball and Dai Suter, and workshop participants through their contributions to discussion on the day of the workshop.

Introduction The aim of the workshop was to review the current knowledge on agronomic potential of herbaceous legumes native to Australia and draw conclusions about the feasibly of domesticating a small number of these plants as grain legume crops for the medium and low rainfall zones of the southern Australian wheatbelt.

Workshop summary There are 14 genera of herbaceous legumes native to arid and semi-arid areas of Australia (Bell et al. 2010). Whilst their agronomic potential was recognised in the 1950s, research has been limited, and has mostly focused on their use as pastures (Kennedia, Cullen, Glycine) or their usefulness as a source of desirable traits for close crop relatives (Glycine) (Britten and Delacy 1979; Millington 1958; Rees et al. 1993; Silsbury 1958). In the last decade there have been a number of small projects which investigated agronomic potential of native herbaceous legumes; most of these projects were linked to the Future Farm Industries CRC and its predecessor, the CRC for Plant-based Management of Dryland Salinity. Small-scale germplasm collections and basic evaluation were undertaken. However, the focus of these activities was perennial species for pastures (Ryan et al. 2008 ).

It was hoped that records of use of native legume seed by aboriginal people could give an indication of species with good potential for human consumption. Unfortunately, the very small amount of literature on this topic suggests that herbaceous legumes were no more than an occasional source of food for aboriginal people (in contrast to some species of wattles and, in some instances, grass seeds) (Brand-Miller and Holt 1998). We can only speculate why. We know that to collect by hand a volume of seeds worth processing (grinding, cooking etc) of some legume species is slow and energy intensive when compared to cereal species, due to factors such as sparse populations of small plants, staggered seed ripening, dropping seeds and small seeds (see Abbo et al. 2008). Some legume seed may also commonly contain toxins. However, many Near Eastern grain legume wild relatives had similar disadvantages and yet were still domesticated in prehistoric times (Abbo et al. 2008). Indeed, perhaps these unfavourable characteristics provided impetus for domestication, i.e. the move towards cultivation of a dense stand, for species that were identified as a desirable food. Overall, the factors that lead to domestication of legumes in the Near East remain poorly understood, but recent work suggests that alongside productivity considerations, food nutritional value, such as the content of favourable amino acids like tryptophan, may have played a role (Abbo et al. 2009). Interestingly in the Near East, domestication tended to select single species from a number of closely related, and apparently similar, taxa. For example, the wild progenitor of chickpea is Cicer reticulatum, while other sympatrically-distributed species in the primary (ie. C. echinospermum) and secondary gene pool of the cultigens were passed over during the domestication process (Ladizinsky and Adler 1976). The reasons for this selective domestication are not known, but may have been nutritional (Abbo et al. 2009). It would be interesting to know if these reasons are still relevant today, in view of current diet and farming systems, as this could suggest whether use or non-use of a species of native legumes by Australian aborigines should be considered when prioritising species for development as grain legumes. Further focused investigation of traditional use of native legumes by aboriginal people is required both to ensure all existing knowledge is recorded and to classify species as “novel food” or not according by

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Food Standards Australia New Zealand (FSANZ) (FSANZ 2010). Significant time and funding is required to show a novel food is safe for human consumption.

The 16 genera that contain native species of herbaceous legumes are Canavalia, Crotalaria, Cullen, Desmodium, Glycine, Glycyrrhiza, Hardenbergia, Indigofera, Kennedia, Lespedeza, Lotus, Rhynchosia, Swainsona, Tephrosia, Trigonella and Vigna (Bell et al. 2010; pers. comm. Richard Snowball). A number of these genera (e.g., Glycine, Crotalaria, Trigonella and Vigna) include already cultivated exotic grain legumes. Using the existing literature, Bell et al. (2010) evaluated species based on the extent that their natural distribution corresponded to arid and semi-arid climatic regions, as well as existing information on traits related to harvestability (uniformity of ripening, propensity to retain pod, pod shattering and growth habit), grain qualities (seed size, chemistry and colour, absence of toxins) and fecundity. The Australian species of Vigna, Canavalia, Tephrosia 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. The species for which sufficient information was available that were considered highest priority for further investigation were Glycine canescens, Cullen tenax, Swainsona canescens, S. colutoides, Trigonella suavissima, Kennedia prorepens, Glycyrrhiza acanthocarpa, Crotalaria cunninghamii and Rhynchosia minima. However, it was noted that published data on seed yield were rare and information on many other traits was also limited. Unpublished data from pasture germplasm nurseries run as part of CRC activities in Western Australian and South Australia, which was presented at the workshop, suggested several native species possess traits desirable in grain legumes, but few have a large number of desirable traits. Accessions from southern locations had most potential due to more suitable flowering times. Overall, the data from Western Australia indicated the following as priority species: 1) Glycine canescens; 2) Swainsona canescens, S. colutoides, S. purpurea and S. swainsonioides; 3) Lotus cruentus; and 4)Kennedia prorepens. The South Australian data suggested Swainsona, Cullen and Kennedia possess most potential, with best performing species again including Swainsona canescens, S. colutoides, S. swainsonioides and C. tenax.

Overall, research to date suggests that there are a number of genera and species of native legumes that show potential to be developed as grain legumes. However, we don’t know if the results obtained are indicative of a species, as most data came from only a small number of accessions (collections); often only one. The variation that exists within wild populations of the species is not known. To assess this variation we are currently constrained by limited ex-situ germplasm collections for most of the genera of interest (e.g. Cullen 166, Kennedia 24, Lotus 41 and Swainsona 108 accessions). Most accessions were collected in South Australia and Western Australia (CRC activities) and there are few from the populations known from herbarium records (CHAH 2010) to occur in Victoria, New South Wales, the Northern Territory and Queensland. Some collections may be available from specimens held in botanic gardens or herbarium collections. These collections have not been assessed by the participants in the workshop. As these collections are not usually stored in a way that preserves the longevity of seed, any seed available is likely to be of poor quality. However, some effort to access uncollected or uncommon species from these sources in future research endeavours may prove worthwhile.

To our knowledge, many species of Cullen, Swainsona and Kennedia have not yet been collected, although we know of their existence and distribution from herbarium records (CHAH 2010). For instance, for Cullen, 11 species have been collected, while 14 are uncollected and hence unavailable for evaluation. In contrast, there are 67 500 accessions of the exotic grain legume chickpea (Cicer arietinum) recorded in genebanks. Thus, increasing germplasm collections is a priority as all future domestication activities may be compromised by lack of diverse germplasm. Moreover, an investment in field collection of germplasm will be very small compared to later costs of a breeding program. A diverse, but small and hence easy to work with, collection could be made of priority species by careful, targeted field collecting of representative accessions. However, we first need to understand the factors that control genetic diversity in field populations. This can only be achieved through collecting widely across the full range of a small number of priority species, followed by determination of the factors (e.g. latitude, longitude, rainfall etc) that correlate with genetic diversity.

For instance, research in Israel on wild emmer wheat (Triticum turgidum ssp. dicoccoides) found greatest genetic diversity and best drought resistance in populations not from areas with extreme aridity, as may be expected, but from areas with intermediate levels of aridity which are exposed to the greatest climatic fluctuations (Peleg et al. 2007). Indeed, in very dry areas, plants may evolve to avoid drought by becoming ephemeral. The findings of Peleg et al. (2007) are yet to be tested in the Australia, but they do suggest that in

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the search for native legumes for the low rainfall cropping zones of Australia, species and accessions from medium and variable rainfall environments should not be excluded.

The above example makes clear the importance of understanding the factors controlling genetic variability. An alternative approach would be to make small targeted collections of germplasm following climate matching for target agricultural regions (Bennett et al. 2010) (i.e. identifying native species which naturally grow in a climate similar to that of a target region). However, it must be noted that this approach may miss suitable germplasm. For instance, subtropical accessions of Cullen performed unexpectedly well during the summer in the Mediterranean climate of the wheatbelt of Western Australia (Bennett et al. 2008). Thus, species or accessions not expected to perform well in an agricultural region based on their climate of origin can be useful in unexpected ways, such as providing green feed in response to summer rainfall (which is likely to increase in parts of southern Australia) or providing good agronomic or disease resistance characteristics that can be transferred using a breeding program to better-adapted types.

The way in which a native grain legume would fit into farming systems can only be broadly speculated upon at the moment. Having originated in soils of poor fertility and in regions with low and variable rainfall, it seems one application may be for low-input farming systems in areas with low or decreasing rainfall. Indeed, recent research, albeit on a small number of species, shows many natives grow well under low phosphorus conditions (Pang et al. 2010a; Pang et al. 2010b). However, this does not provide a justification for a sole focus on natives and some of the novel exotic species examined in the same study also grew very well under low phosphorus conditions. Interestingly, some natives may have a superior ability to access poorly soluble soil phosphorus sources using root exudates; a potentially useful trait in the face of the rapid decline in world phosphorus sources. Whilst many native plants are known to develop symptoms of toxicity at very low rates of phosphorus addition, this is not the case for herbaceous legumes, with the exception of Kennedia species, and thus they will grow well in soils in our current agricultural systems (Pang et al. 2010a; Pang et al. 2010b). Superior drought tolerance has been found in some native legumes (Suriyagoda et al. 2010), although unpublished work again suggests that some novel exotic perennial herbaceous legumes also posses this trait (pers. comm. Jiayin Pang). More thorough evaluation of individual species and genera is required to determine how native legumes may best fit agronomically into farming systems. There seems to be no reason why native grain legumes could not also be developed for conventional high input farming systems in medium rainfall areas.

Development of a native grain legume must be closely guided by consultation with the food industry. When developing a new product for human use there are a large number of factors to consider. These include the importance of matching consumer trends (e.g. currently, wholegrain) but also finding a point of difference to existing options (e.g. “native”, “sustainable”, “healthy”). A nation-wide supply of a reasonable tonnage in the 100s – 1000s of tonnes that is consistent year to year is desirable and growing areas must be close to infrastructure in order to reduce freight costs. A minimum of 12 months shelf life is required and flexibility of use is desirable (i.e. bread, health bars, breakfast cereals etc). There are many other requirements which include an ability to flow easily in and out of a silo. It can be difficult to support a price premium and thus yield is very important. For use as stockfeed it is especially important to have consistency of supply and nutrient value. For a native legume product destined for use by either humans or livestock, detailed compositional data would be required.

Some useful lessons may be taken from the recent domestication in Western Australia of the widely grown exotic annual grain legume Lupinus angustifolius (narrow-leafed lupin). Many unfavourable characters, which are also present in native legumes and in the wild relatives of other major exotic grain legumes in the Middle East, have been relatively easily addressed in the domestication process. Traits that have been incorporated include soft-seededness to allow easy germination, two non-shattering genes to ensure seed doesn’t fall onto the ground at maturity, low seed alkaloids to make seed suitable for human and livestock consumption, white seed, and reduced time to flowering to allow adaptation to environments with short growing seasons (Gladstones 1994). Subsequently breeding efforts have increased yield responsiveness, harvest index (the proportion of biomass allocated to grain) and fecundity (pers. comm. Jens Berger), resulting in yield increases at around 2.5% per year (Stefanova and Buirchell 2010). Early uptake by progressive farmers was noted as an important factor in the success of narrow-leafed lupin. Overall, successful wide-scale adoption of narrow- leafed lupin followed three decades of activities based around a dedicated breeding and evaluation program (Nelson and Hawthorne 2000). A strong message from this case-study is the crucial importance of public

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funding to the maintenance of a research program of sufficient size to achieve such an impressive outcome so quickly.

Recommendations It was concluded by workshop participants that further evaluation of herbaceous native legumes with the aim of development of a grain legume crop is warranted. For the genera containing most of the priority species listed above, i.e. Swainsona, Cullen and Kennedia, further work should commence with collection of seed and evaluation of plant characters for species not yet collected. Simultaneously, for a small subset of species already known to show considerable promise (e.g. Glycine canescens, Swainsona canescens, S. colutoides, Kennedia prorepens and Cullen tenax), collection trips should be undertaken with the aim of assembling a collection of sufficient size and diversity to allow determination of which factors control genetic diversity. This new knowledge will be crucial to then guide development of diverse, but small, germplasm collections for priority species. Efforts should be made to involve aboriginal people in collecting activities and to tap into any unrecorded knowledge on herbaceous native legumes. The history of lupin domestication in Australia suggests the activities recommended above are unlikely to be funded by private breeding companies and, therefore, would need to be largely funded by public money; as would subsequent breeding and selection programs, and the development of agronomic packages. In addition, consultation and collaboration with representatives of the food industries that would purchase a native grain legume product must also occur at all stages of the domestication process to ensure the final product is well-suited to industry needs.

References Abbo S, Saranga Y, Peleg Z, Kerem Z, Lev-Yadun S, Gopher A (2009) Reconsidering domestication of legumes versus cereals in the ancient Near East. Quarterly Review of Biology 84, 29-50. Abbo S, Zezak I, Schwartz E, Lev-Yadun S, Kerem Z, Gopher A (2008) Wild lentil and chickpea harvest in Israel: bearing on the origins of Near Eastern farming. Journal of Archaeological Science 35, 3172-3177. Bell L, Bennett R, Ryan M, Clarke C (2010) The potential of herbaceous native Australian legumes as grain crops: a review. Renewable Agriculture and Food Systems, in press. Bennett RG, Colmer TD, Real D, Ryan MH (2008) New perennial pasture legumes: persistence and productivity of Australian Cullen species on deep acid sands in WA’s low-rainfall wheatbelt. In '2nd International Salinity Forum'. 31st March to 3rd April, Adelaide Convention Centre, Adelaide. Bennett RG, Ryan MH, Colmer TD, Real D (2010) Prioritisation of novel pasture species for use in water-limited agriculture: a case study of Cullen in the Western Australian wheatbelt. Genetic Resources and Crop Evolution, in press. Brand-Miller JC, Holt SHA (1998) Australian Aboriginal plant foods: a consideration of their nutritional composition and health implications. Nutrition Research Reviews 11, 5-23. Britten E, Delacy I (1979) Assessment of the genetic potential for pasture purposes of the Psoralea eriantha- patens complex, a native legume of the semi-arid zone Australian Journal of Experimental Agriculture 19, 53-58. CHAH (2010) Australia's virtual herbarium. Council of Heads of Australasian Herbaria. http://www.ersa.edu.au/avh/about.jsp. Accessed 01/08/2010. FSANZ (2010). Food Standards Australia New Zealand. http://www.foodstandards.gov.au/consumerinformation/novelfoods/. Accessed 01/08/2010. Gladstones JS (1994) An historical review of lupins in Australia. In '1st Lupin Technical Symposium'. Perth, WA pp. 1-38. (Department of Agriculture). Ladizinsky G, Adler A (1976) Genetic relationships among the annual species of Cicer L. Theoretical and Applied Genetics 48, 197-203. Millington AJ (1958) The potential of some native West Australian plants as pasture species. Journal of the Royal Society of Western Australia 42, 1-6. Nelson P, Hawthorne WA (2000) Development of lupins as a crop in Australia. In: Linking Research and Marketing Opportunities for Pulses in the 21st Century. In 'Proc. 3rd Int. Food Legumes Res. Conf.' (R. Knight, Ed.), Kluwer Academic Publishers, Dordrecht, pp. 549-559. Pang J, Tibbett M, Denton MD, Lambers H, Siddique KHM, Bolland MDA, Revell CK, Ryan MH (2010a) Variation in seedling growth of 11 perennial legumes in response to phosphorus supply. Plant and Soil 328, 133- 143. Pang J, Ryan MH, Tibbett M, Cawthray GR, Siddique KHM, Bolland MDA, Denton MD, Lambers H (2010b) Variation in morphological and physiological parameters in herbaceous perennial legumes in response to phosphorus supply. Plant and Soil 331, 241-255.

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Peleg Z, Fahima T, Saranga Y (2007) Drought resistance in wild emmer wheat: physiology, ecology, and genetics. Israel Journal of Plant Sciences 55, 289-296. Rees M, Jones R, Brown A (1993) Glycine latifolia – a potentially useful native legume for clay soils in tropical and subtropical Australia. In 'Proceedings of the XVII International Grassland Congress'. Rockhamption, QLD, Australia. Ryan M, Bennett R, et al. (2008 ) Searching for native perennial legumes with pasture potential. In '14th Australian Agronomy Conference, 21-25 September 2008'. Adelaide, South Australia. (Ed. M Unkovich). (Australian Society of Agronomy.). Silsbury JH (1958) Agricultural potentialities of the genus Kennedya Vent. in Western Australia. The Journal of the Australian Institute of Agricultural Science 24, 337-338. Stefanova KT, Buirchell B (2010) Multiplicative mixed models for genetic gain assessment in lupin breeding. Crop Science 50, 880-891. Suriyagoda LDB, Ryan MH, Renton M, Lambers H (2010) Multiple adaptive responses of Australian native perennial legumes with pasture potential to grow in phosphorus- and moisture-limited environments. Annals of Botany 105, 755-767.

75 Native Legumes as a Grain Crop for Diversification in Australia

by Megan Ryan, Lindsay Bell, Richard Bennett, Margaret Collins and Heather Clarke

Publication No. 10/223

This report is the first comprehensive assessment of the potential It will also be of interest Australia-wide to research funding for native Australian herbaceous legumes to be developed as bodies, scientists, plant breeders, innovative growers and grain legume crops for the drier areas of the Australian grain members of the food industry. belt. A review of the literature, along with assessment of various sources of unpublished information, was used to prioritise RIRDC is a partnership between government and industry species. Grain quality and plant growth characteristics relevant to invest in R&D for more productive and sustainable rural to domestication potential, especially harvestability, were then industries. We invest in new and emerging rural industries, a assessed in a glasshouse experiment for 17 species of native suite of established rural industries and national rural issues. legume. Three commercial grain legume cultivars were included for comparison. Most of the information we produce can be downloaded for free or purchased from our website . This report is targeted at people interested in using our native flora to improve the sustainability of southern Australian RIRDC books can also be purchased by phoning cropping systems, especially in the context of a drying climate. 1300 634 313 for a local call fee.

Front cover photo: Seed pods of Glycine canescens are produced in the midst of the herbaceous canopy Back cover photo: The twining habit displayed by these Glycine canescens plants is common among progenitors of many modern crop legumes

Contact RIRDC: Level 2 15 National Circuit Ph: 02 6271 4100 Most RIRDC publications can be viewed and purchased at Barton ACT 2600 Fax: 02 6271 4199 our website: Email: [email protected] PO Box 4776 web: www.rirdc.gov.au www.rirdc.gov.au Kingston ACT 2604 Bookshop: 1300 634 313

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