Perennial Grain Crops for High Water Use The case for Microlaena stipoides

A report for the Rural Industries Research and Development Corporation by CL Davies, DL Waugh and EC Lefroy February 2005 RIRDC Publication No 05/024 RIRDC Project No UWA 60A

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© 2005 Rural Industries Research and Development Corporation. All rights reserved. ISBN 1 74151 123 2 ISSN 1440-6845 Perennial grain crops for high water use Publication No. 05/024 Project No. UWA 60A 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 industries. The information should not be relied upon for the purpose of a particular matter. Specialist and/or appropriate legal advice should be obtained before any action or decision is taken on the basis of any material in this document. The Commonwealth of Australia, Rural Industries Research and Development Corporation, the authors or contributors do not assume liability of any kind whatsoever resulting from any person's use or reliance upon the content of this document. This publication is copyright. However, RIRDC encourages wide dissemination of its research, providing the Corporation is clearly acknowledged. For any other enquiries concerning reproduction, contact the Publications Manager on phone 02 6272 3186. Researcher Contact Details Dr Ted Lefroy CSIRO Sustainable Ecosystems Private Bag 5, PO Wembley WA 6913 Phone: 08 9333 6442 Fax: 08 9333 6444 Email: Ted.Lefroy@cse.csiro.au

Dr Christine Davies CLIMA/The University of Western Australia 35 Stirling Highway Crawley WA 6009 Phone: 08 6488 1432 Fax: 08 6488 1140 Email: chrisdav@cyllene.uwa.edu.au

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 1, AMA House 42 Macquarie Street BARTON ACT 2600 PO Box 4776 KINGSTON ACT 2604 Phone: 02 6272 4819 Fax: 02 6272 5877 Email: rirdc@rirdc.gov.au. Website: http://www.rirdc.gov.au Published in February 2005 Printed on environmentally friendly paper by Canprint

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Foreword This research set out to domesticate an Australian native grass to produce a perennial grain crop. Perennial (long-lived) grain crops offer a new solution to the long-standing problems of salinity and soil erosion associated with conventional cropping systems based on annual plants. Perennial grain crops have two potential advantages over annual crops in the sustainable management of soils and landscapes. They do not require annual cultivation and, being perennial, they make more complete use of annual rainfall. The starting point for this project was the observation that there are two forms of perennial plant; woody and herbaceous, and within each of these there are two potential end products; vegetative (leaves and stems) or reproductive (seeds and fruit), giving four possible production systems. Three of these already exist in Australia as forestry (woody and vegetative), perennial pastures (herbaceous and vegetative) and horticulture (woody and reproductive). The system not currently exploited commercially is perennial herbaceous plants grown for grain harvest (herbaceous and reproductive). The team identified the native species weeping rice grass (Microlaena stipoides) as a promising candidate for a perennial grain crop on the basis of its seed size and wide distribution. This project was funded from RIRDC Core Funds which are provided by the Australian Government. This report, an addition to RIRDC’s diverse range of over 1200 research publications, forms part of our Resilient Agricultural Systems R&D program, which aims to foster the development of agri-industry systems that have sufficient diversity, integration, flexibility and robustness to be resilient enough to respond opportunistically to continued change. Most of our publications are available for viewing, downloading or purchasing online through our website: • downloads at www.rirdc.gov.au/fullreports/index.html • purchases at www.rirdc.gov.au/eshop Tony Byrne Acting Managing Director Rural Industries Research and Development Corporation

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Acknowledgments

This work was jointly funded by RIRDC, GRDC, The University of Western Australia and CSIRO. Wal Whalley (University of New England, Armidale), Meredith Mitchell (Department of Primary Industries, Rutherglen) and Ian Chivers (Native Seeds Pty Ltd, Melbourne) provided advice and support throughout the project. Michelle Murphy (UNE, Armidale) collected microlaena for this study in NSW and Meredith Mitchell assisted in the collection in WA. Jens Berger assisted with the statistical analysis.

Glossary of Terms

Culm an aerial stalk or stem of grain and grasses that bears flowers

Spikelet a unit of the inflorescence consisting of one or more florets, the rachilla and the glumes

Floret in grasses the unit composed of a lemma and palea and the small flower they enclose

Rachilla the main stem of a grass spikelet above the glumes

Glumes the outer bract of a spikelet

Husk spikelet minus the seed, including infertile florets, palea and lemma surrounding the seed, and the rachilla

Palea in a grass floret, the upper one of the two bracts enclosing a flower

Lemma the lower of two bracts enclosing a grass flower

Seed naked caryopsis

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Contents Foreword ...................................................................................................................................iii Acknowledgments..................................................................................................................... iv Glossary of Terms ..................................................................................................................... iv Executive Summary .................................................................................................................. vi Introduction ................................................................................................................................ 1 Materials and Methods............................................................................................................... 5

Germplasm Collection......................................................................................................................... 5 Site and preparation............................................................................................................................. 6 Measurements...................................................................................................................................... 6

Seed yield ........................................................................................................................................ 7 Culm and spikelet number............................................................................................................... 8

Statistical analysis ............................................................................................................................... 8 Results ........................................................................................................................................ 9

NSW collection ................................................................................................................................... 9 WA collection ................................................................................................................................... 10 Correlations with seed yield .............................................................................................................. 11 Principal Components Analysis ........................................................................................................ 11

Discussion ................................................................................................................................ 13 Future work ....................................................................................................................................... 15

Implications and Recommendations ........................................................................................ 16 References ................................................................................................................................ 19 Appendix 1. Annotated Bibliography ..................................................................................... 21 Appendix 1. Annotated Bibliography ..................................................................................... 21

A. Australian perennial grasses with potential for domestication ..................................................... 21 B. Exotic perennial grasses with potential for use as a human food ................................................. 29 C. Further detail on Microlaena stipoides ......................................................................................... 32

Appendix 2. Perennial grass species with potential for domestication................................... 39 A. Native species............................................................................................................................... 39 B. Exotic species ............................................................................................................................... 40

Appendix 3. Networks............................................................................................................. 42

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Executive Summary One of the big challenges facing Australian agriculture is the management of salinity. The problem is that the annual, high seed–yielding crop plants like wheat and barley that are the backbone of our agriculture make incomplete use of annual rainfall. Excess rainfall accumulates in the soil and mobilises ancient stores of salt. Australia’s native vegetation on the other hand is well adapted to ancient soils and highly variable rainfall. Our woodlands and heath are dominated by plants that are long lived and use all the rainfall. The downside is that they grow more slowly than annual plants and produce less harvestable material each year. The challenge is to find commercially valuable perennial plants. This research investigated the potential to domesticate an Australian native grass (Microlaena stipoides) to produce a perennial grain crop. Perennial grain crops offer a new solution to the long-standing problems of salinity and soil erosion associated with conventional cropping systems based on annual plants.

Seed yield and its components (culm number, spikelet number per culm, seed set and seed weight) were measured in 46 accessions of Microlaena stipoides (microlaena, meadow or weeping rice grass) from Western Australia and New South Wales to quantify potentially useful variation in the species.

A high degree of variability was found to exist, with a twenty-fold range in seed yield (0.1 to 2.4 g/plant), five-fold range in seed weight (129 to 666 mg per 100 seeds), two-fold range in spikelet number (14 to 30 per culm), eight-fold range in seed set (12 to 98%) and a five-fold range in culm number (11 to 59 per plant). Seed yield was positively and significantly (P<0.05) correlated with culm number, seed set and seed weight (r>0.55 for all). No correlation was found between seed yield and spikelet number per culm (r=-0.14).

The range in seed yield and its components suggests there is sufficient variation within microlaena to make selections for higher yielding lines. This variation will enable breeders to exploit genetic diversity more efficiently and identify useful accessions for further work. This study evaluated the variation in components of seed yield at only one site. The next step should be to evaluate superior lines at multiple sites across Australia.

While it is unlikely that grain yields in microlaena will be comparable with those of annual grains, it is possible that a dual value as a grain and grazing crop could help to offset lower returns from seed harvest. Other favourable characteristics such as synchronous maturity and resistance to shattering need to be considered.

Continued consideration of Microlaena stipoides as an alternative perennial grain crop requires evaluation of the compositional and nutritional characteristics of its grain to determine its suitability for use in food products for human consumption and stock feed. Of particular interest are its starch composition and, because it is fairly closely related to rice, its likely gluten-free status.

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Introduction One of the big challenges facing Australian agriculture is the management of dryland salinity. The problem is that high seed–yielding annual crop plants such as wheat and barley, the backbone of our farming systems, make incomplete use of annual rainfall. Excess rainfall accumulates in the soil and mobilises ancient stores of salt. Australia’s native vegetation on the other hand is well adapted to ancient soils and highly variable rainfall. Our woodlands and heath are dominated by plants that are long lived and use virtually all the rainfall (Specht 1999). The disadvantage with basing agriculture on perennial plants is that they grow more slowly than annual plants and produce less harvestable material each year. The challenge is to find native perennial plants with the potential for domestication. The starting point for this project was the observation that there are two forms of perennial plant that could potentially be used in agricultural systems; woody and herbaceous, and within each of these there are two potential end products; vegetative (leaves and stems) or reproductive (seeds and fruit), giving four possible production systems. Three of these already exist in Australia in the form of forestry (woody and vegetative), perennial pastures (herbaceous and vegetative) and horticulture (woody and reproductive). The system not currently exploited commercially is perennial herbaceous plants grown for grain harvest (herbaceous and reproductive). The concept of perennial grain crops presents major biological and agronomic challenges. The biological challenge is that perennial crops appear to contradict the fundamental principle of biology that plants partition their resources either towards reproduction or perenniality, but not both. The agronomic challenge is demonstrating an exception to a 10 000 year-old rule that successful grain production is based on fitting high seed-yielding annual grasses into a short seasonal window to avoid climatic extremes (Ewel 1999; Jackson and Jackson 2000). A review of perennial grain crop research produced two notable conclusions: 1) long life and high seed yield are not necessarily mutually exclusive, with some experimental perennial crops yielding in excess of 1 t/ha of grain in the USA (Tripsacum dactyloides (Jackson and Dewald 1994)), and 2) there are likely to be niches in agricultural landscapes where low grain yield from a perennial cereal crop could be complemented by its grazing value (Wagoner 1990a). Evidence that perenniality and high seed yield are not mutually exclusive comes from recent research in the USA on one of the progenitors of corn Tripsacum dactyloides (eastern gamagrass or sesame grass) which showed that high seed yield (>1 t/ha) in a pistillate mutant of this perennial grass did not compromise its perenniality (Jackson and Dewald 1994; Piper 1998). Additional evidence comes from a perennial wheat breeding program at the University of California in Davis in the 1940s that produced intergeneric hybrids between wheat (Triticum aestivum) and a perennial relative Agropyron ponticum. Lines bred specifically for perennial habit yielded within 70% of the best commercial wheats of their time (Suneson 1959; Suneson and Pope 1946). The suggestion that there is a new niche in agricultural landscapes for multi-purpose perennial grain and grazing plants comes from research in Europe and North America. Intergeneric hybrids of Triticum and Agropyron developed by the United States Department of Agriculture in the 1930s (Sando 1935; Vinall and Hein 1937) are undergoing selection at Washington State University in Pullman with the aim of developing perennial grain and grazing crops as an alternative to annual grain crops on hilly erodible land (Cox et al. 2002; Scheinost et al. 2001); a concept first put forward by (Jackson 1985). Jackson and co-workers at The Land Institute in Kansas are developing three perennial crops; two cereals (Tripsacum dactyloides and Leymus racemosus, (Jackson and Jackson 2000) and a perennial grain legume (Desmanthus illinoisensis) which is yielding in excess of 1 t/ha (Kulakow et al. 2000). Following contact with that group, the principal investigator of this project Dr Ted Lefroy was invited to become a member of The Land Institute’s Advisory Board.

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In Hungary, a perennial rye Gyakorlati agroforum has been released for grain and forage production (Hodosne-Kotvics et al. 1999), while at Utah State University, Indian ricegrass (Oryzopsis hymenoides) is under development (Jones et al. 1998; Jones and Nielson 1992) and in south west USA, selections of perennial saltgrass (Distichis spicata) are reported to have grain yields of 0.5 t/ha (J Leake pers comm). The two perennial grain crops with a history of cultivation are Thinopyron intermedium and Leymus arenarius. Thinopyron intermedium or intermediate wheatgrass appears to have been harvested for human consumption in Turkey, Armenia and the Caucasus mountains during Byzantine times (Wagoner 1990ab, 1994, 1995b; Wagoner and Schauer 1990). The cultivar Triga has been released in the United States as a multi-purpose grain and grazing plant. Leymus arenarius is being studied as a potential grain crop in Iceland where it has been used as a food grain since the time of the Vikings (Anamthawat-Jonsson 1996; Griffin and Rowlett 1981). The trade-off with a perennial grain crop is that grain yields will not be as high as that of annuals. It is hoped that their dual role as grain and grazing plants will help to offset lower returns from seed harvest. In this way it is planned to produce a plant tailored to a particular niche, the arable mid to upper slopes of the medium to high rainfall zones where there has been an increase in the area under cropping in recent years in response to declining wool prices, but where recharge and soil erosion represent significant risks to sustainable land use. This niche is estimated to cover some 3–5 million hectares across the Australian grain belt. Under dual-purpose perennial grain crops, this land could remain commercially viable and environmentally sustainable. The original objective of this project was to screen perennial relatives of the major grain crops (wheat, barley, rye), plus high seed-yielding naturalised and endemic perennial grasses for their potential as grain crops. The outcome would be perennial grass accessions ranked on the basis of seed yield, seed size, seed head architecture, harvest index and rooting depth as an indicator of their potential for further development as high water use perennial grain crops. Three classes of perennial grasses were selected as target groups: 1. Perennial relatives of the major grain crops (wheat, barley, rye) in the tribe Triticeae which share

their basic genome with the annual crops. Specific examples include accessions of Thinopyron intermedium from which the US grain cultivars were selected, Secale montanum, S. strictum, Hordeum spontaneum and H. bulbosum.

2. Perennial grasses endemic to southern Australia, seed of which was used for human consumption by aboriginal people (eg. Astrebla lappacea, A. pectinata and A. squarosa, Eragrostis spp.).

3. Relatives of exotic summer-active, drought-tolerant perennial grasses that have become naturalised across the cropping zone, mostly from southern Africa (eg. Eragrostis spp., Chloris spp. Ehrharta spp.).

A literature review of endemic and exotic perennial grasses with potential for domestication and/or grain production was completed (Appendices 1 and 2). We searched for evidence of the use of Australian native grasses for food by aboriginal people and the only evidence we found was with annual plants grown in the region shown in Figure 1. In order to collect additional information on the use of perennial grasses in Australia, both exotic and endemic, we contacted research teams, non-government organisations (NGOs) and community groups either interested in or currently working on perennial grasses as a means of surveying current work. A list of these groups and contact details is contained in Appendix 3. As a result of discussions with this network of researchers and producers, and our reading of the literature on introduced perennial grasses (eg. Londsdale 1994), we decided that the environmental weed risk associated with exotic naturalised plants and perennial relatives of the major grain crops was too great and consequently we concentrated on perennial grasses endemic to southern Australia.

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Figure 1. Distribution of the aboriginal and contemporary grain belts in Australia indicating the location of specific studies listed in Appendix 1 (1Allen 1974; 2Cane 1989; 3Cleland and Johnston 1936; 4Cleland and Johnston 1939a; 5Cleland and Johnston 1939b; 6Maggiore 1985; 7O’Connell 1983). As a consequence of our literature review on native perennial grasses and our contact with networks, we identified the Australian native species Microlaena stipoides (microlaena, meadow or weeping rice grass) as a promising candidate on the basis of seed size (Figure 2) and wide distribution (Figure 3). Microlaena has large seeds, approaching that of commercial rice varieties, with larger examples being 5–6 mm long and weighing 4–6 mg (Whalley and Jones 1997). It also has indeterminate flowering in that many genotypes will continue to produce seed throughout the summer, provided soil water is available (Earl 1993). Microlaena has a wide geographic distribution throughout southern Australia particularly in the coastal and tableland regions of eastern Australia and the medium to high rainfall areas of southern and western Australia. It tolerates acid soils, drought and frost, and is capable of year round green growth with high forage digestibility (55–80%) and quality (10–27% crude protein) (Waters et al. 2000). Seed yield is an essential criterion for the commercial acceptability of both grain crops and perennial pasture species. It is determined by four main reproductive features — seed set, seed weight, number of reproductive culms and number of spikelets per culm (Wagoner 1990a). Seed production in microlaena compares very favourably with introduced perennial grass species and far exceeds documented yields of other native grass species (Earl et al. 1994). Measuring the variation in components of seed yield amongst germplasm accessions collected across Australia will enable breeders to exploit genetic diversity and identify useful accessions more efficiently. We expect that the variation in seed yield and its components in Microlaena stipoides will be sufficient to identify superior accessions for the development of a perennial grain crop.

aboriginal grain belt (after Tindale 1974)

contemporary grain belt

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4

2

3

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Figure 2. Seed of a NSW line of Microlaena stipoides (left) shown with long grain white rice.

Figure 3. Distribution of Microlaena stipoides in Australia (Australian Virtual Herbarium 2004).

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Materials and Methods Germplasm Collection Seed collection of Microlaena stipoides from naturally occurring stands was carried out in northern New South Wales and southern Queensland in April/May 2001, targeting areas where large-seeded accessions had been previously reported by a University of New England research team (RDB Whalley, pers comm). A total of 34 accessions, comprised of seed from multiple plants at each location, were collected (Figure 4, inset 1) and are referred to as the NSW collection. In June 2001, seedlings were propagated in jiffy pots containing a 50:50 peat:coarse sand mix with Osmocote®. Seedlings were reared in a glasshouse and received regular overhead watering. In September 2001, 1020 seedlings from the NSW collection were transplanted into the field at the UWA Field Station at Shenton Park. The plants were arranged in a randomised block design with three replicate plots consisting of ten plants. Seedlings were widely spaced in double rows 350 mm apart on 600 mm wide plastic matting with 350 mm spacing between plants in a row. The rows of plastic matting were spaced 800 mm apart. A collecting trip in the south west of Western Australia was carried out in December 2001, targeting areas where microlaena had been reported in the WA Herbarium’s database (FloraBase). Seedlings were collected with roots intact, placed in plastic bags and kept moist. The bags were sealed to prevent moisture loss. A total of 12 accessions, comprised of 30–40 plants from each location, were collected (Figure 4, inset 2) and are referred to as the WA collection. Seedlings were transplanted directly into 200 mm pots containing potting mix (equal quantities of white sand, peat, pine bark) with Osmocote®. The seedlings were kept in the shade house and received regular overhead watering. In May 2002, 360 of the original seedlings collected from WA were established in the field plots at Shenton Park (as per the NSW collection).

Figure 4. Collection sites of Microlaena stipoides germplasm in Western Australia and New South Wales/southern Queensland.

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Site and preparation The UWA Field Station is located in Shenton Park, approximately 8 km west of Perth’s CBD (31° 56’ 55” S, 115° 47’ 33” E). The site, located under bird netting, contains virgin Karrakatta sands of the Spearwood dune system with a pH of 8.0. Before sowing the soil was fertilised with 300 kg/ha Super Potash 3:2 and sprayed with 225 l/ha metham sodium and 2.5 l/ha Roundup® for weed control. The soil was then rotary-hoed to a depth of about 200 mm. Plastic sheeting (600 mm wide) was laid for weed suppression. Plants were watered with overhead sprinklers as required, usually every second or third day during the warmer months (September to the break of season). All plants received applications of Urea or NPK at a rate equivalent to 50 kg N/ha every three months. A weed control program consisted of hand weeding individual plants and rotary hoeing between rows as required. Measurements In the 2001/02 season (NSW collection only) plants were assessed for spikelet yield, seed weight, culm number and weight. Seed yield was determined by collecting spikelets over an 8-week period from 17 December 2001 to 12 February 2002. In the 2002/03 season (NSW and WA) plants were assessed for spikelet yield, seed weight, seed set, culm number, length and weight and spikelet number per culm. Seed yield was determined by collecting spikelets over an 8-week period from 23 October 2002 to 18 December 2002. In the 2003/04 season plants were assessed for spikelet yield, habit and plant height. Seed yield for the NSW collection was determined by collecting spikelets over a 6-week period from 21 November 2003 to 5 January 2004 and for the WA collection over a 4-week period from 1 November 2003 to 2 December 2003. We noted that plants from the NSW collection flowered twice each year with seed production at the end of the year and mid-year (not measured) while those in the WA collection flowered only once with seed collection at the end of the year. The four components of seed yield examined were seed set, seed size, number of culms per plant and number of spikelets per culm (Wagoner 1990). Measurements of spikelet yield, 100-seed weight, culm number and spikelet number per culm were collected at the replicate level meaning that within accession variance was not assessed. Caution must be exercised when making comparisons between seed yield and its components of the two collections as the NSW collection was in its second year of production while the WA collection was transplanted from the field and assumed to be in its first year of production. For the purposes of this study a spikelet consists of three florets, one fertile and two infertile, and the rachilla (stalk). The fertile floret potentially holds one caryopsis (seed). The ‘husk’ is the spikelet minus the seed, that is, the two infertile florets, the palea and lemma surrounding the seed, and the rachilla (Figure 5).

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Figure 5. Line drawing of Microlaena stipoides (modified from Gardner 1952) identifying habit, culm, spikelet, pedicel, glumes, rachilla, two awned infertile lemmas, palea and lemma surrounding fertile seed. Seed yield Spikelets were collected by hand over the harvest period by gently clasping individual tillers bearing mature spikelets and running the hand along the inflorescence. Care was taken not to remove immature spikelets which were allowed to ripen for harvest at a later date. On each collection date spikelets from each replicate were placed into paper envelopes and allowed to air dry. After a period of at least two weeks the spikelets were weighed. Weights from each collection date were added together for a total spikelet weight per replicate. In the 2002/03 season, one hundred spikelets from each replicate were selected at random and weighed. The number of empty spikelets were separated, counted and weighed. The remaining spikelets (filled) were separated into seeds and husks; each was weighed. This data enabled an estimation of the number of seeds set as a percentage of spikelet number and seed yield (Table 1).

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Table 1. Method for calculating seed yield and seed set of Microlaena stipoides in the 2002/03 season.

Description Formula A 100 Spikelet weight (g) as measured B Number empty spikelets per 100 spikelets as measured C Weight empty spikelets per 100 spikelets (g) as measured D Seed set (%) 100–B E Weight of filled spikelets per 100 spikelets (g) as measured F Seed weight per 100 spikelets (g) as measured G 100 seed weight (g) (F/D)*100 H Spikelet yield per plant (g) as measured I Filled spikelet yield per plant (g) H–((H/A)*C) J Seed yield per plant (g) (I*F)/E

Culm and spikelet number At the end of the seed collecting periods in 2001/02 and 2002/03, when all the spikelets had been removed from the plant, the culms were removed and collected by cutting them off 30 mm above the base of the plant (ground level). One hundred culms from each replicate were randomly selected from the total; the number of spikelets per culm was determined for 50 of these culms by counting the pairs of glumes persistent on the rachis. The 100-culms were tied together to keep them separate from the total and then oven-dried at 60°C for 48 h. Total culms and 100-culms were weighed. The number of culms per plant was estimated [((total culm weight/100-culm weight) x 100)/number of plants]. Plant height and habit In 2003/04 plant height and habit (erect, semi-erect, semi-prostrate, prostrate) were recorded. Statistical analysis The data were analysed using Genstat for Windows 6.1. Following tests of normality, analyses of variance (ANOVA) were calculated on all the variables as one-way ANOVAs to identify variation among accessions within a trait within a season and two-way ANOVAs to identify variation between years. Correlations were performed on all variables to identify relationships among traits. For the 2002/03 season, principal components analysis (PCA) was performed on the means of seed yield and its components (seed weight, seed set, culm number, spikelet number) for each accession of individual collections. PCA was performed on the correlation matrix rather than the covariance matrix as the variables were measured on different scales. PCA defines new variables, known as principal components, each of which is a linear combination of the original variables. The principal components are uncorrelated with each other, with the first principal component explaining as much of the variation as possible, the second principal component explaining as much of the remaining variation, and so on. The maximum number of principal components that can be defined is the same as the original number of variables. However, most of the variation in the data can often be accounted for by the first few principal components. Graphical presentation of the accession means of the first and second components for each accession allows identification of groupings and patterns among the accessions. Arrows showing the directions of the original variables in relation to the principal components (as indicated by the latent vectors, also known as loadings) can be presented on the same axes, enabling further interpretation of the accession means.

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Results NSW collection The range of means and grand means of measured traits of the NSW collection in the 2001/02, 2002/03 and 2003/04 seasons are shown in Table 2. There was significant (P<0.05) variation among the 34 accessions for most of the characters measured when averaged over the three replicates in any one season. Seed yield differed between accessions within a season with at least a 10-fold range (Table 2). The highest seed yield of 2.39 g/plant was produced by LIG183 in 2002/03. There was no difference (P>0.05) in seed yield between seasons. Mean seed yield was 0.75 g/plant in 2001/02, 1.42 g/plant in 2002/03 and 0.83 g/plant in 2003/04. One hundred seed weight had a three-fold range from 129 mg (MM013) to 359 mg (LIG183) with a grand mean of 290 mg. Accessions varied in their ability to set seed, ranging from 12 to 98 seeds per one hundred spikelets (mean 84). The proportion of seed weight in spikelet yield had a seven-fold range from 9 to 63% (mean 54%). Culm length differed among accessions (P<0.001) but there was no difference in culm weight or culm number per plant. The number of spikelets per culm differed between accessions (P<0.001) ranging from 14 to 30 (mean 20). There was no difference in plant height between accessions. Plant habit ranged from semi-erect to semi-prostrate. Table 2. The range of means and the grand mean of seed yield and its components in the NSW collection (n=34, 3 reps) of Microlaena stipoides grown at the Shenton Park Field Station from 2001 to 2004. NSW 2001/02 2002/03 2003/04

Range of means

Grand mean

Range of means

Grand mean

Range of means

Grand mean

Spikelet yield (g/plant) 0.63–2.22 1.37 ** 1.02–4.25 2.55 ns 0.58–2.72 1.52 ***

Seed yield (g/plant) 0.15–1.42 # 0.75 *** 0.13–2.39 1.42 *** 0.12–1.52 # 0.83 ***

100 seed weight (mg) 129–359 290 *** Seed weight per spikelet yield (%) 8.8–63.1 53.9 ***

Seed set (%) 12.0–98.0 83.8 *** Spikelet number/culm 14.1–30.3 20.1 *** Culm number/plant 10–62 38 ** 20–59 40 ns Culm length (mm) 493–789 667 *** Culm weight (g/plant) 1.57–8.88 4.58 *** 3.56–13.84 8.44 ns Plant height (mm) 317–550 446 ns

Habit (score 1–4) ## 2.5–3.5 3.1 * # Seed yield calculated from ratio determined in 2002/03 season. ## Habit: 1 = erect, 2 = semi-erect (45–90°), 3 = semi-prostrate (0–45°), 4 = prostrate Significant differences between accessions * P<0.05, ** P<0.01, *** P<0.001, ns not significant.

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WA collection The range of means and grand means of measured traits of the WA collection in the 2002/03 and 2003/04 seasons are summarised in Table 3. Wide variation was observed (P<0.05) among the 12 accessions for most of the characters investigated when averaged over the three replicates in any one season. Seed yield varied between accessions within a season: in 2002/03 it ranged from 0.41 g/plant (Mitchell River) to 2.40 g/plant (Stretch) with a grand mean of 1.39 g/plant; in 2003/04 it ranged from 0.19 g/plant (Mitchell River) to 1.48 g/plant (Hacks) with a grand mean of 0.99 g/plant (Table 3). There was no difference (P>0.05) in seed yield between seasons. One hundred seed weight in 2002/03 had a two-fold range with the smallest being 299 mg (Mitchell River) and the largest 666 mg (Tenterden) with a grand mean of 494 mg. In 2003/04 100-seed weight ranged from 412 mg to 789 mg with a mean of 579 mg. Seed weight did not differ (P>0.05) between seasons. Accessions varied in their ability to set seed, ranging from 28 to 81 seeds per one hundred spikelets (mean 68). The proportion of seed weight in spikelet yield had a three-fold range from 15 to 48% (mean 41%). Culm length differed among accessions (P<0.05) but there was no difference in culm weight, culm number or spikelet number per culm. Plant height ranged from 350 mm to 500 mm with a mean of 425 mm. Plant habit ranged from semi-erect to semi-prostrate. Table 3. The range of means and the grand mean of seed yield and its components in the WA collection (n=12, 3 reps) of Microlaena stipoides grown at the Shenton Park Field Station from 2002 to 2004. WA 2002/03 2003/04

Range of means Grand mean Range of means Grand mean

Spikelet yield (g/plant) 1.81–5.12 3.27 ns 1.24–3.45 2.32 ***

Seed yield (g/plant) 0.41–2.40 1.39 ** 0.19–1.48 # 0.99 ***

100 seed weight (mg) 299–666 494 ** 412–789 579 *

Seed weight per spikelet yield (%) 15.1–48.3 41.0 *** Seed set (%) 27.7–81.0 68.3 *** Spikelet number/culm 17.7–24.4 19.8 ns Culm number/plant 11–27 20 ns Culm length (mm) 697–917 814 * Culm weight (g/plant) 4.57–9.75 8.00 ns Plant height (mm) 350–500 425 ***

Habit (score 1–4) ## 2.0–3.0 2.3 * # Seed yield calculated from ratio determined in 2002/03 season. ## Habit: 1 = erect, 2 = semi-erect (45–90°), 3 = semi-prostrate (0–45°), 4 = prostrate Significant differences between accessions * P<0.05, ** P<0.01, *** P<0.001, ns not significant.

11

Correlations with seed yield The results of correlation analyses between seed yield and all other measured traits for each collection in each year are shown in Table 4. The correlation matrices revealed correlations between seed yield and many of the variables. A strong positive correlation exists between seed yield and spikelet yield in both the NSW and WA collections in all seasons (r = 0.86–0.97). Other positive correlations with seed yield were seed weight (r = 0.61–0.68), seed weight per spikelet yield (r = 0.67, 0.75), seed set (r = 0.69, 0.77), culm number (r = 0.80, NSW only), culm length (r = 0.49, NSW only), culm weight (r = 0.67–0.79) and plant habit (r = 0.70, WA only). There was no relationship (P>0.05) between seed yield and spikelet number per culm or plant height in either collection. Table 4. Correlation (r values) between seed yield and all other measured traits for the WA and NSW collection in each year. NSW Collection WA Collection

Trait 2001/02 2002/03 2003/04 2002/03 2003/04

Spikelet yield (g/plant) 0.86 ** 0.91 ** 0.88 ** 0.92 ** 0.97 **

100 seed weight (mg) 0.65 ** 0.68 * 0.61 *

Seed weight per spikelet yield (%) 0.67 ** 0.75 **

Seed set (%) 0.69 ** 0.77 **

Spikelet number/culm –0.15 –0.14

Culm number/plant 0.81 ** 0.80 ** 0.55

Culm length (mm) 0.49 ** 0.21

Culm weight (g/plant) 0.79 ** 0.70 ** 0.67 *

Plant height (mm) 0.26 –0.02

Habit (score 1–4) # –0.18 0.70 * Significant differences * P<0.05, ** P<0.01, ns not significant. # Habit: 1 = erect, 2 = 45–90°, 3 = 0–45°, 4 = prostrate Principal Components Analysis The principal components analysis revealed that principal component 1 (PC1) explained 65% of the variation present in the NSW collection (Figure 6a) and 54% in the WA collection (Figure 6b) while principal component 2 (PC2) explained 19% of the variation present in both collections. The variables with the highest loadings on PC1 were seed weight, seed yield and seed set and, in the NSW collection, culm number. PC2 had a high loading for spikelet number per culm and, in the WA collection, culm number. In the NSW collection, the accessions with the highest seed yield and seed size combinations were LIG183, MM004, MM028, MM005 and MM009 (Figure 6a) and in the WA collection they were Stretch, Tenterden, Kent River, Unicup, Porongorups and Young’s Siding (Figure 6b). Seed yield of MM004 was driven by high culm numbers while that of MM009 was driven by high seed weights and seed set. Seed yield of LIG183, MM005 and MM028 was associated with high seed weights, seed set and culm numbers. Seed yield of Porongorups was driven by high culm numbers while that of Kent River was driven by high seed weights. Seed yield of Porongorups, Unicup, Tenterden and Stretch were associated with high seed weights, seed set and culm numbers.

12

-7

-5

-3

-1

1

3

5

7

-7 -5 -3 -1 1 3 5 7

Seed yield

(a)

PC1: 64.7%

PC2: 18.8%

Seed w eight

Culm number

Spikelet number

Seed set

LIG 183MM009

MM004

MM005 MM028

-4

-3

-2

-1

0

1

2

3

4

-4 -3 -2 -1 0 1 2 3 4

Seed yield

(b)

PC1: 54.0%

PC2: 18.7%

Seed w eight

Culm number

Spikelet number

Seed set

Young's Siding

UnicupPorongorups

Tenterden

Stretch

Kent River

Figure 6. Principal components analysis of seed yield and its components of Microlaena stipoides from (a) the NSW collection (34 accessions) and (b) the WA collection (12 accessions). The arrows on each biplot represent the loadings on each principal component for each of the characters scored (ie. the latent vectors). The accessions with the most desirable traits are named and marked ( ).

13

Discussion A high degree of variability was found to exist within seed yield and its components of Microlaena stipoides, with a twenty-fold range in seed yield, five-fold range in seed weight, two-fold range in spikelet number per culm, eight-fold range in seed set and a five-fold range in culm number. This range suggests there is sufficient variation within microlaena to make selections for higher yielding lines. There was no discernable relationship between seed yield or its components and location or habitat type. The range in variation probably reflects its complex breeding system. Understanding the breeding system of any grass species is critical before starting a breeding or selection program (Smith and Whalley 2002). The majority of microlaena inflorescences are produced with cleistogamous spikelets (fertilisation occurs within closed flowers) and a minority with chasmogamous spikelets (fertilisation occurs in open flowers) (Groves and Whalley 2002). As a result, microlaena tends to be uniform within accessions and highly variable between them which relates to the high degree of inbreeding that occurs in this species (Clifford 1962; Groves and Whalley 2002). Aerial cleistogamous florets in microlaena usually have two very tiny anthers, each producing relatively few pollen grains (Groves and Whalley 2002). The pollen grains usually germinate within the anthers and the pollen tubes grow out through the anther wall and onto the stigmatic surface (Clifford 1962; Connor and Matthews 1977). This form of pollination is common to many cleistogamous species (Uphof 1938). Pollination usually occurs while the spikelets are still enclosed within the sheath of the flag leaf. As a result, emasculation and crossing would present severe practical difficulties. Chasmogamous florets, on the other hand, usually have four large anthers with lots of pollen allowing and fertilisation usually occurs well after the spikelets emerge. Therefore, it should be relatively easy to emasculate florets and make crosses. However, it is first necessary to learn how to reliably stimulate the production of chasmogamous inflorescences in microlaena before crosses can be made routinely (Huxtable 1990). The multiplication of resultant selected genotypes would be easy using only seed produced from cleistogamous inflorescences. Seed yield is an important criterion in determining the commercial acceptability of new perennial grass cultivars. It is a major limitation to the domestication and adoption of many native grasses with most being best described as conservative seed producers, partitioning photosynthate to maximise plant survival rather than seed production (Smith and Smith 1997; Wagoner 1990). This trade-off means that grain yields from a perennial are unlikely to be as high as that from an annual crop (Wagoner 1990). Encouraging evidence that challenges this view is the work of Jackson and Dewald (1994) who showed that high seed yield (>1 t/ha) in a pistillate mutant of Tripsacum dactyloides (eastern gamagrass) did not compromise its perenniality. Where perennial grasses are used for multiple uses, their forage value and lower input costs compensate for lower grain yields. High seed yield is a significant characteristic for a perennial grass whether it is used for grain or fodder production. Reproductive yield of native grasses is often expressed as spikelet yield rather than seed yield. For the purposes of grain production in this study, the true seed yield of microlaena was determined, which ranged from 9 to 63 per cent of spikelet yield. In the literature on perennial grass pasture research, spikelet yield is often taken to be synonymous with seed yield. In any research aimed at grain yield for human or animal consumption, seed yield should refer to the weight of the naked caryopses. Seed production of Microlaena stipoides compares favourably with that of introduced perennial grass species and exceeds documented yields of other Australian native grass species (Earl et al. 1994). In this study, seed yields ranged from 0.4 to 2.4 g/plant which equates to 11 to 196 kg/ha (48 to 418 kg/ha of spikelets) at the plant densities sown. These yields per hectare are necessarily low as they were obtained from small plots with a density of only 8 plants/m2. Assuming that yield per plant does

14

not decrease with plant density up to 26 plants/m² (Pukeridge and Donald 1967; Wade et al. 1992), this would be equivalent to seed yields in the range 35 to 550 kg/ha. Our results are supported by the work of Cole et al. (2001) who reported seed yields in the microlaena accession LIG 183 of 132, 287 and 503 kg/ha over three successive years. Whalley and Jones (1997) achieved spikelet yields of up to 700 kg/ha from 225 m² plots by hand and vacuum harvesting on a weekly basis over a four month period. Assuming the ratio of seed to spikelets is 0.54 (grand mean from our NSW collection) seed yield would be in the order of 380 kg/ha. Very high spikelet yields in three lines from NSW were reported by Earl (1993) ranging from 34 to 72 g/plant; assuming the above seed yield to spikelet yield ratio of 0.54, seed yields would range from 18 to 39 g/plant. These high yields were partly due to a long harvest period of six months. While the grain yield of microlaena is unlikely to be as high as that of an annual crop — the national average wheat yield in Australia has ranged from 1140 to 2100 kg/ha over the last decade (Australian Wheat Board Ltd 2003) — it compares favourably with the yields of other native and exotic perennial grasses. Spikelet yields of lines of native grasses selected for accelerated development in the LIGULE program (Cole et al. 2001) ranged from 31 to 226 kg/ha in wallaby grass (Austrodanthonia fulva), 41 to 239 kg/ha in kangaroo grass (Themeda triandra), 42 to 139 kg/ha in red grass (Bothriochloa macra), 151 to 518 kg/ka in tall windmill grass (Chloris ventricosa), 44 to 376 kg/ha in common wheatgrass (Elymus scaber), 28 to 62 kg/ha in curly windmill grass (Enteropogon acicularis). Loch et al. (1996) measured spikelet yields of up to 200 kg/ha in curly Mitchell grass (Astrebla lappacea). Seed yields in the native North American switchgrass (Panicum virgatum) ranged from 20 to 98 kg/ha (Boe 2003) up to 450 kg/ha (Vogel and Moore 1993). Studies in the United States of grain yields in intermediate wheatgrass (Thinopyrum intermedium) were generally the highest in their first year of production (100–600 kg/ha) (Wagoner 1995; Wagoner and Schauer 1990). Some accessions with high seed weight, seed set and seed yield during their first year of production continued to exhibit good yield potential in subsequent years, while others dropped dramatically (Wagoner 1995). Selection of superior lines should therefore be based on those maintaining a high seed yield over a longer period. The components of seed yield are strongly influenced by an interaction of genetic, environmental and crop management factors. The WA collection had heavier seeds (299–666 mg per 100 seeds) than the NSW collection (129–359 mg per 100 seeds). One hundred seed weights of the three accessions studied by Earl (1993) ranged from 290 to 330 mg. Waters et al. (2000) reported the released pasture cultivar Shannon having a seed weight of 500 mg per 100 seeds. These weights compare favourably with the established perennial grain crop, intermediate wheatgrass, where one hundred seed weights ranged from 350 to 820 mg (Wagoner 1995; Wagoner and Schauer 1990). Seed set is defined here as the percentage of spikelets containing a seed. Seed set was higher in the NSW collection with an average of 84 per cent (range 12–98%) than in the WA collection with an average of 68 per cent (range 28–81%). Seed set ranged from 12 to 59 per cent in common wheatgrass (Elymus scaber) (Murphy and Jones 1999) and 3 to 66 per cent in intermediate wheatgrass (Wagoner 1995). In other work, seed set has been described as the proportion of seed weight in spikelet yield (Earl 1993). In this study, seed set by weight ranged from 9 to 63 per cent (mean 51%). This is supported by the work of Earl (1993) who found seed weight contributed 52 to 70 per cent of the total spikelet weight, although these values are likely to be an overestimation as spikelet yield did not include the weight of empty spikelets. In Earl’s study the accession with the highest spikelet weight had the lowest seed set and the one with the lowest spikelet weight had the highest seed set resulting in almost the same seed yield for both accessions. There was no such relationship in our research. The number of spikelets per culm is often used as an indicator of yield potential (Griffiths et al. 1980) however in our case seed yield was not related to the number of spikelets per culm. High seed yields in

15

microlaena were largely associated with high culm numbers in both collections. Among the three microlaena accessions studied by Earl (1993) the component contributing most to seed yield varied between seed size, culm number or spikelet number per culm. Consistent among those three accessions was the large amount of variance in total yield accounted for by culm number (74–91%). It is likely that increased seed production in microlaena is largely dependent on the production of a high number of reproductive culms, particularly in the NSW collection. Selection for high culm number earlier in development rather than at harvest may increase the rate of selection of higher seed yielding lines. Future work This study evaluated the variation in components of seed yield at one site. The next step should be to evaluate superior lines, identified in Figure 6, at multiple sites across Australia. These lines (except for MM009, MM028 and Young’s Siding) were established in 30 m² plots at the WA Department of Agriculture’s Medina Research Station, Perth in January 2004 for seed increase. While it is unlikely that grain yields in microlaena will be comparable with those of annual grains, it is possible that a dual value as a grain and grazing crop could help to offset lower returns from seed harvest. Other favourable characteristics such as synchronous maturity and resistance to shattering need to be considered. Continued consideration of Microlaena stipoides as an alternative perennial grain crop requires evaluation of the compositional and nutritional characteristics of its grain to determine its suitability for use in food products for human consumption and stock feed. Of particular interest are its starch composition and, because it is fairly closely related to rice, its likely gluten-free status.

16

Implications and Recommendations From this research we identified that there is sufficient variation within Microlaena stipoides to make selections for higher yielding lines. A number of studies exist for further assessment of the potential of Microlaena stipoides as an economically viable dual purpose grain and grazing crop. Examples of these studies are outlined below and graphically presented in a flow chart (Figure 7). 1. This project has identified that a high degree of variability exists within desirable characteristics of

our current germplasm base of 46 accessions. However the extent to how this captures the best properties for microlaena for development as a perennial grain crop is not known. Further collections from a wider range of environments across Australia would broaden the range of genotypes available for selection of high seed-yielding lines with desirable properties.

2. Assess the potential of microlaena as a dual purpose grain and grazing crop in the medium to high

rainfall zones of Australia through data collection at a range of field sites across southern Australia. 3. Agronomic trials in targeted environments across Australia would explore how to optimise grain

yield. Examples of studies include the response to fertiliser and recharge control potential. Native grasses such as microlaena have never been selected for their response to fertilisers. Does microlaena respond to fertiliser applications (eg. N and K)? Does seed and/or biomass yield increase with N and/or K applications?

4. There is evidence to indicate that microlaena has a high protein (21.6%) and a unique amino acid

profile. Nutritional properties of the seed need to be assessed to determine its potential as a human food. Useful nutritional properties would be gluten, starch, total dietary fibre, fat, sugars, protein, anti-nutritional factors (lectins, CTIA, TIA, tannins), moisture and ash contents.

5. The processing properties of microlaena grain needs to be assessed in a range of food applications

such as: a. A whole grain softened and added to products such as breads, muffins, crackers and sweet

biscuits. b. A ground whole grain to be used as a wholemeal flour supplement in breads, biscuits, muffins

and Asian noodle products. c. A flour product to be included in breads, biscuits, muffins and Asian noodle products.

6. If nutritional and processing properties prove promising the next step would be to identify potential

food industry partners for assessment of microlaena as a food product. The pathway to domestication of wild plants is lengthy and other techniques such as molecular markers may need to be used to speed up the process to understanding the genetic relationships by describing genotypes more rapidly and more accurately.

17

Q1.

Doe

s the

seed

of M

icro

laen

a st

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des h

ave

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tiona

l and

pr

oces

sing

pro

perti

es su

itabl

e as

a fu

nctio

nal f

ood

addi

tive?

Yes

N

o

Q2.

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com

mer

cial

ly v

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e to

pro

duce

M.

stip

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s see

d as

a fo

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dditi

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Q3.

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s the

seed

of M

. stip

oide

s hav

e va

lue

as a

who

le g

rain

food

?

Yes

N

o

Com

mer

cial

an

d N

RM

ou

tcom

es

achi

eved

Yes

N

o

Q4.

Is se

ed p

rodu

ctio

n of

M. s

tipoi

des

com

mer

cial

ly v

iabl

e as

a w

hole

gra

in fo

od?

Q5.

Are

hig

h se

ed-y

ield

ing

lines

of M

. st

ipoi

dess

uita

ble

as p

astu

re c

ultiv

ars?

Yes

N

o

Com

mer

cial

an

d N

RM

ou

tcom

es

achi

eved

End

of

Proj

ect

Yes

N

o

NR

M

outc

omes

ac

hiev

ed

End

of

Proj

ect

Crit

erio

n 1

Crit

erio

n 2

Crit

erio

n 3

Crit

erio

n 4

Crit

erio

n 5

Cri

teri

a 2

& 4

•

Prod

uctio

n co

sts

• H

arve

st c

osts

•

Seed

yie

ld

• M

arke

t res

pons

e

Cri

teri

on 5

•

Hig

h se

ed y

ield

•

Veg

etat

ive

yiel

d •

Dig

estib

ility

•

Nut

ritiv

e va

lue

• Ea

se o

f har

vest

•

Low

cos

t see

d

Cri

teri

on 3

•

Glu

ten

• St

arch

& S

ugar

s •

Tota

l die

tary

fibr

e •

Fats

•

Prot

eins

and

am

ino

acid

s •

Ant

i-nut

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nal f

acto

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• Ea

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para

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•

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• St

arch

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l die

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e •

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•

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ino

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nal f

acto

rs

• Fo

od a

pplic

atio

ns

•Fl

our q

ualit

yG

rain

pro

pert

ies

Gra

in p

rope

rtie

s A

gron

omy

Agr

onom

y G

razi

ng p

oten

tial

Res

ults

from

fiel

d sc

reen

ing

of

exis

ting

lines

(U

WA

60A

) and

sc

reen

ing

of n

ew

colle

ctio

ns

Out

puts

and

dec

isio

n po

ints

in th

e co

mm

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al d

evel

opm

ent o

f Mic

rola

ena

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oide

s Fi

gure

7.

18

One avenue for more rapid progress into the development of perennial crops could well be made by looking at crosses between domesticated annual crops and their perennial relatives but this comes with a risk of weediness and root disease. Further investment into the development of perennial crops by the GRDC and CRC for Plant-based Management of Dryland Salinity is subject to a symposium being held at The University of Western Australia in late September 2004. The symposium features two international speakers: Dr Stan Cox, plant breeder at The Land Institute in Salina, Kansas and previously plant breeder with the USDA; and Doug Lammer from Washington State University. The Land Institute is a private not for profit organisation that has been breeding and selecting perennial grain crops over the last 20 years. The WSU team is developing perennial wheats for the Palouse region of the Pacific Northwest. RIRDC has been invited to this symposium and a copy of the findings will be forwarded to the Environment and Farm Management Program.

19

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potential. Buvusindi 10, 101–113. Australian Virtual Herbarium (2004) [Online]. Available at: http://avh.calm.wa.gov.au/cgi-bin/avh.cgi.

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http://www.awb.com.au/AWBL/Launch/Site/AboutAWB/Content/CommunityEducation/GrainProduction. (verified 7 July 2004)

Boe A (2003) Genetic and environmental effects on seed weight and seed yield in switchgrass. Crop Science 43, 63–67.

Clifford HT (1962) Cleistogamy in Microlaena stipoides (Labill.) R.Br. Department of Botany Papers (University of Queensland) 14, 63–72.

Cole I, Johnston W, Metcalfe J, Tooth I, Mitchell M, Koen T (2001) Overcoming the seed production barriers with native grasses [Online]. Available at:

http://www.regional.org.au/au/asa/2001/p/8/cole.htm. (verified 9 July 2004) Connor H, Matthews B (1977) Breeding systems in New Zealand grasses. 7. Cleistogamy in

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Jackson W (2002) Breeding perennial grain crops. Critical Reviews in Plant Sciences 21, 59–91. Earl JM (1993) Seed production of selected accessions of Microlaena stipoides (Labill.) R. Br.

Honours thesis, University of New England, Armidale, New South Wales. Earl JM, Whalley RDB, Jones CE (1994) Variation in the components of seed yield and germination

requirements among three accessions of Microlaena stipoides. In 'Working papers from the 8th biennial rangeland society conference'. Katherine, Northern Territory. pp. 179–180. (Australian Rangeland Society: Alice Springs)

Ewel JJ (1999) Natural systems as models for the design of sustainable systems of land use. Agroforestry Systems 45, 1–21.

Gardner CA (1952) 'Flora of Western Australia. Gramineae. Volume 1.' (Government Printer: Perth) Griffiths DJ, Lewis J, Bean EW (1980) Problems of breeding for seed production in grasses. In 'Seed

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157–182. Hodosne-Kotvics G, Krisztian J, Dornbach L (1999) Perennial rye: a novel variety released from a

new interspecific hybrid rye. Gyakorlati Agroforum 10, 63. Huxtable CHA (1990) Ecological and embryological studies of Microlaena stipoides (Labill.) R.Br.

Dissertation thesis, University of New England. Jackson L, Dewald C (1994) Predicting evolutionary consequences of greater reproductive effort in

Tripsacum dactyloides, a perennial grass. Ecology 75, 627–641. Jackson W, Jackson LL (2000) Developing high seed yielding perennial polycultures as a mimic of

mid-grass prairie. In 'Agriculture as a Mimic of Natural Ecosystems'. (Eds EC Lefroy, RJ Hobbs, MH O'Connor, JS Pate) pp. 1–55. (Kluwer: Dordrecht)

Jones TA, Majerus ME, Scheetz JG, Holzworth LK, Nielson DC (1998) Registration of 'Rimrock' Indian ricegrass. Crop Science 38, 539–540.

Jones TA, Nielson DC (1992) High seed retention of Indian ricegrass PI478833. Journal of Rangeland Management 45, 72–74.

20

Kulakow PA, Benson LL, Vail JG (1990) Prospects for domesticating Illinois bundleflower. In 'Advances in new crops: proceedings of the first national symposium — new crops, research, development, economics'. Indianapolis. (Eds J Janick, JE Simon). pp. 168–171. (Timber Press: Portland, Oregon)

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Murphy MA, Jones CE (1999) Observations on the genus Elymus (Poaceae: Triticeae) in Australia. Australian Systematic Botany 12, 593–604.

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Uphof JCT (1938) Cleistogamous flowers. Botanical Review 4, 21–49. Vogel K, Moore K (1993) Native North American grasses. In 'New crops'. (Eds J Janick, JE Simon)

pp. 284–293. (John Wiley and Sons, Inc: New York) Wade LJ, Douglas ACL, Bell KL (1992) Effect of plant density on grain yield and yield stability of

diverse grain sorghum hybrids. AIAS Occasional Publication 68, 414–422. Wagoner P (1990) Perennial grain development: past efforts and potential for the future. Critical

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Wagoner P, Schauer A (1990) Intermediate wheatgrass as a perennial grain crop. In 'Advances in new crops: proceedings of the first national symposium — new crops, research, development, economics'. Indianapolis. (Eds J Janick, JE Simon) pp. 143–145. (Timber Press: Portland, Oregon, USA)

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Whalley RDB, Jones CE (1997) 'Commercialising the Australian native grass M. stipoides.' RIRDC Publication No 97/34 (Rural Industries Research and Development Corporation: Canberra)

21

Appendix 1. Annotated Bibliography A. Australian perennial grasses with potential for domestication 1. Abele K (1959) Cytological studies on the genus

Danthonia. Transactions of the Royal Society of South Australia 82, 163–173.

2. Aldous D, Chivers I (1996) Agronomic and quality

factors of weeping grass (Microlaena stipoides). New Zealand Turf Management Journal February, 31–32. Scope: agronomic and quality factors of six accessions of weeping grass Perennial grass species: Microlaena stipoides Detail: agronomic characters: no tillers, no. rhizomes, leaf length to width, leaf width, culm length. Quality characters: uniformity, density, texture, growth habit, smoothness and colour. There was considerable variation between accessions.

3. Allen H (1974) The Bagundji of the Darling Basin:

cereal gatherers in an uncertain environment. World Archaeology 5, 309–322. Scope: Seed gathering activities of the Bagundji Aboriginal population in the western Murray Darling Basin, NSW Perennial grass species: Panicum decompositum (mentioned here as an annual) – seeds between December and March; Panicum effusum (perennial?) – grew abundantly on the flooded lands near creeks; Panicum miliaceum (perennial?) Detail: collecting methods

4. Allan C (1997) Factors affecting landholder opinion

of Microlaena stipoides (Labill.) R.Br. Masters thesis, University of New England, Armidale.

5. Anon (1992) Danthonia richardsonii and D. linkii.

Plant Varieties Journal (Australia) 5(1), 18–21. 6. Anon. (1995) Variety: 'Griffin' syn. '703.6.12'.

Application no. 95/052. Plant Varieties Journal 8, 22. 7. Anon. (1995) Variety: 'Shannon' syn. '17.2.6.5.12'.

Application no. 94/124. Plant Varieties Journal 8, 22. 8. Anon. (1995) Variety: 'Wakefield' syn. '39.1.8.2.5'.

Application no. 94/125. Plant Varieties Journal 8, 22. Scope: agronomic characteristics Perennial grass species: Microlaena stipoides Detail: cvs. Griffin, Shannon, Wakefield and two ecotypes. Plant height, growth habit, leaf colour, leaf attitude, flag leaf width & length, inflorescence length, number of spikelets/inflorescence

9. Baxter A (Ed.) (2000) 'Native grasses information kit.'

(Agriculture Western Australia: Perth). Scope: Useful native grasses in south west WA Perennial grass species: 60 species mentioned of which 12 are mentioned to occur in the wheatbelt area (Amphipogon caricinus, Aristida contorta, Aristida holathera, Aurostipa juncifolia, Elymus scaber,

Hemarthria uncinata, Microlaena stipoides, Monachather paradoxus, Poa porphyroclados, Tetrarrhena laevis, Themeda triandra, Thryidolepis multiculmis). Detail: perennial native grasses in WA (no suggested uses for grain production), seed sources, list of information sources and consultants, list of references, websites, species information sheets (Themeda triandra, Microlaena stipoides), seed establishment, harvesting techniques, several useful papers (eastern states) in kit. Microlaena stipoides – seed is produced in Nov/Dec and, in response to summer and autumn rains, from January to May. Remains green throughout the year. More commonly seen in damp or semi-shade areas. Major component of Mount Lofty Ranges woodlands. 30–70 cm. High frost and drought tolerance. As many as 100 viable cuttings can be obtained from large plants by simple cutting the rhizomic root system at each root node. Seed size looks to be 10 mm x 1.5 mm. Seed is obtained by stripping florets off the long narrow panicle at the end of the weeping seed stalk. As much time as practicable should be allowed for the seed to mature before harvesting. When pinched between the thumbnail and forefinger viable seed will feel stiff and firm in comparison to the hollow and easily bent infertile florets. A good average crop should yield 10 g dry seed/m2 with 70–80% viability. Themeda triandra – can be distinguished by the red/brown tinge on the older leaves and its rusty red coloured ripe seed heads on stems with dark nodes. 40–90 cm high. Moderate frost tolerance. High drought tolerance.

10. Beardsell D (1985) Domestication problems of

Australian plants. In 'The food potential of seeds from Australian native plants. Proceedings of a colloquium held at Deakin University on 7 March 1984'. (Ed GP Jones) pp. 147–159. (Deakin University Press: Victoria)

11. Beckers DJ (1993) Seed and seedling establishment

biology of Microlaena stipoides (Labill.) R.Br. BSc(Hons), University of New England, Armidale, NSW.

12. Brand JC, Cherikoff V (1985) Nutrients in native

plant seeds — 1. In 'The food potential of seeds from Australian native plants Proceedings of a colloquium held at Deakin University on 7 March 1984'. (Ed. GP Jones) pp. 31–45. (Deakin University Press: Victoria). Scope: Nutrient contents of seeds of various shrubs and grasses that Aborigines used for food in the desert regions of Central Australia Perennial grass species: Eragrostis eriopoda Detail: Eragrostis eriopoda – eaten as a raw paste, 17.4% protein, high in iron (31 mg/100 g)

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13. Brock RD, Brown JAM (1961) Cytotaxonomy of

Australian Danthonia. Australian Journal of Botany 9, 62–91.

14. Cane S (1989) Australian Aboriginal seed grinding

and its archaeological record: a case study from the Western desert. In 'Foraging and Farming: the evolution of plant exploitation'. (Eds D Harris, G Hillman) pp. 99–119. (Unwin Hyman: London). Scope: Distribution, use and importance of seeds; technology used to process seeds; the extent and nature of the probable correlation between the archaeological record and the relative importance of seeds to Aboriginal people who lived in the region (Great Sandy Desert) Perennial grass species: 15 grass species (7 perennial?) –Brachiaria miliiformis, Dactyloctenium radulans, Eragrostis eriopoda, Eragrostis laniflora, Panicum australiense, Panicum cymbiforme (Whiteochloa cymbiformis), Panicum decompositum Detail: Panicum australiense – responds quickly to fire, grows in dense concentrations, produces an abundance of seeds, 13% protein Panicum cymbiforme – 8.5% protein

15. Cashmore AB (1932) An investigation of the

taxonomic and agricultural characters of the Danthonia group. Bulletin No. 69 CS&IR.

16. Chivers IH, Aldous DE (1996) Response of weeping

grass (Microlaena stipoides) to pre-emergence and post-emergence herbicides. In 'Proceedings of the 3rd ATRI Turf Research Conference'. Brisbane, Qld & Sydney NSW pp. 80–87. Scope: studies were conducted with weeping grass to investigate the phytotoxicity caused by trifluralin (as Treflan), dithiopyr (as Dimension), pendimethalin (as Stomp), bensulide (as Exporsan), 2, 4-D as the sodium salt formulation (as Yates Lawn Weed and Feed) and MCPA and dicamba as the dimethylamine salt formulation (as Hortico Clover & Bindii). Perennial grass species: Microlaena stipoides Detail: pre-emergence spring applications of pendimethalin, at rates of 2.0 and 4.0 l/ha, and autumn applications of trifluralin at 1.5 l/ha, were effective on germinating weedy growth, with limited phytotoxic effects recorded on the emerging weeping grass seedlings. MCPA and dicamba was significantly more effective in controlling white clover growth and flower production than the 2,4-D formulation. Neither formulation had any long term phytotoxic effect on the weeping grass.

17. Cleland JB, Johnston TH (1936) Notes on native

names and uses of plants in the Musgrave Ranges region. Oceania 8, 208–215; 328–342. Scope: details of native plants in the Musgrave Ranges – information obtained on an expedition accompanied by two tribal men.

Perennial grass species: Eragrostis eriopoda, Themeda triandra Detail: very little detail on grasses

18. Cleland JB, Johnston TH (1939) Aboriginal names

and uses of plants in the Northern Flinders Ranges. Transactions of the Royal Society of South Australia 63, 172–179. Scope: details of native plants in the Northern Flinders Ranges – information obtained on an expedition accompanied by tribal members. Perennial grass species: Astrebla pectinata, Panicum decompositum Detail: Panicum decompositum – seeds used. Astrebla pectinata – seeds ground and eaten.

19. Cleland JB, Johnston TH (1939) Aboriginal names

and uses of plants at the Granites, Central Australia. Transactions of the Royal Society of South Australia 63, 22–26. Scope: details of native plants in the Granites, ~ 600 km north west of Alice Springs – information obtained on an expedition. Perennial grass species: Themeda triandra, Astrebla spp., Panicum decompositum Detail: very little detail on grasses

20. Clifford HT (1962) Cleistogamy in Microlaena

stipoides (Labill.) R.Br. University of Queensland, Department of Botany Papers 14, 63–72.

21. Cochrane D (1998) Native perennial grasses for

Western Australia. Land Management Society Newsletter November, 11–12. Scope: identifies native perennial grasses with a degree of agricultural or amenity benefit in south west Western Australia (no mention of grain production) Perennial grass species: Microlaena stipoides, Danthonia occidentalis, Danthonia selacea, Neurachne alopecuroides, Amphibromus needsii, Stipa pycnostachya

22. Cole I, Dawson I, Mortlock W, Winder S (2000)

'Guidelines 9: using native grass seed in revegetation.' (Florabank: Canberra)

23. Connor H, Matthews B (1977) Breeding systems in

New Zealand grasses. 7. Cleistogamy in Microlaena. New Zealand Journal of Botany 15, 531–534. Perennial grass species: Microlaena stipoides Detail: single, clandestine, cleistogamic spikelets occur in leaf axils (additional to the previously reported cleistogamic and chasmogamic systems in aerial inflorescences).

24. Crane CF, Carman JG (1987) Mechanisms of

apomixis in Elymus rectisetus from eastern Australia and New Zealand. American Journal of Botany 74, 477–496.

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25. Cribb AB, Cribb JW (1981) 'Useful wild plants of Australia.' (Collins: Sydney). Scope: Australian native grasses – grazing, outlines distribution of 12 native grass species (5 perennial). Perennial grass species: Astrebla spp. (A. lappacea & A. pectinata), Danthonia spp., Dichanthium sericeum, Eragrostis eriopoda, Panicum decompositum, Themeda australis (Themeda triandra) Detail: Astrebla spp. – central west of Qld across the NT into WA, southward into north western part of NSW. A. pectinata most useful in WA, drought resistant (250–500 mm rainfall), dense tussocks, seed heads are spikelike, similar in form to wheat. Danthonia spp. – tussocky, fine-leaved, about 30 cm tall, temperate grasslands and woodlands, some species in all states. Dichanthium sericeum – drier woodland areas in north eastern Australia, tufted grass with hairy spikelets borne on a cluster of 2–4 spiked, each grain has a long erect awn, when not in flower or seed the grass can often be recognised by the tuft of long white hairs at the nodes of the stem, frost sensitive. Eragrostis eriopoda – semi arid areas of inland Australia, drought tolerant, deep roots, small-seeded spikelets, poor fodder but important in drought. Panicum decompositum – warmer parts of Australia, frost sensitive, dies back during winter but develops rapidly from its perennial rootstock following the early summer rains, freely branched seed heads with numerous small seeds. Themeda australis – widespread from the tropical wet summer forests of northern Australia to the temperate woodlands of Victoria and Tasmania, it has mostly gone from the temperate woodlands of southern Australia including the south west of WA, erect, tufted, when mature it has a yellow or reddish colour, with short, broad, whiskery clusters of spikelets produced at intervals along the flowering stem, readily eaten by stock.

26. Earl JM (1993) Seed production of selected

accessions of Microlaena stipoides (Labill.) R.Br. BRurSci(Hons) dissertation, University of New England.

27. Earl JM, Whalley RDB, Jones CE (1994) Variation in

the components of seed yield and germination requirements among three accessions of Microlaena stipoides. In 'Eighth Biennial Rangeland Society Conference'. Katherine, Northern Territory pp. 179–180. (Australian Rangeland Society, Alice Springs). Scope: Seed production and components of seed yield of three accessions of Microlaena stipoides were measured over one growing season Perennial grass species: Microlaena stipoides Detail: Final seed yield did not differ among accessions although there were differences in partitioning. The accession with the greatest number of inflorescences had the least number of spikelets/inflorescence, wile the accession with the least inflorescences/plant had the most spikelets/inflorescence. Seed yields ranged from 1.7–2.2 t/ha.

28. Garden DL, Dowling PM (19??) Native grass-based

pastures. Scientific Research Series TB47. 29. Gott B (1985) The use of seeds by Victorian

Aborigines. In 'The food potential of seeds from Australian native plants Proceedings of a colloquium held at Deakin University on 7 March 1984'. (Ed. GP Jones) pp. 25–30. (Deakin University Press: Victoria). Scope: Seeds of native plants used for food by Victorian Aborigines – not many were used. Perennial grass species: Panicum prolutum, Panicum effusum, Panicum decompositum (mentioned here as annual or perennial)

30. Grice AC, Bowman A, Toole I (1995) Effects of

temperature and age on the germination of naked caryopses of indigenous grasses of western New South Wales. The Rangeland Journal 17, 128–137. Scope: effects of temperature and age on the germination of native caryopses of nine native grasses from western NSW. Detail: details of seed testing. A wide range of characteristics must be considered when assessing the potential of native grasses for domestication. Important amongst them are the characteristics of the seed, germination requirements and the effects of seed storage conditions on seed survival and germination (Lodge and Groves 1991). Domestication requires that large amounts of seed can be easily produced, harvested and processed and that seed retains high viability during storage. Touches on germination and storage – naked caryopses vs. diaspores

31. Groves RH, Hagon MW, Ramakrishnan PS (1982)

Dormancy and germination of eight populations of Themeda australis. Australian Journal of Botany 30, 373–386.

32. Groves RH, Whalley RDB (2002) Grass and

grassland ecology in Australia. Flora of Australia 43, 157–182. Perennial grass species: Dicanthium, Eragrostis, Sporobolus, Elymus, Themeda, Microlaena etc. Detail: Some features of the seed of some major Australian grass genera; an assessment of the ecological significance of the breeding systems shown by representative genera; distribution of the major grassland types occurring in Australia in terms of the climatic regions that they inhabit, their floristics at the generic level and their functional ecology with respect to various environmental factors, including changes in land use; conservation status of Australian grasslands, especially in relation to their invasion by introduced species.

33. Hagon MW (1976) Germination and dormancy of

Themeda australis, Danthonia spp., Stipa bigeniculata, and Bothriochloa macra. Australian Journal of Botany 24, 319–327.

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34. Hodgkinson K (1996) ‘The life and times of perennial grasses.’ http://www.dwe.csiro.au/research/rangelnd/perennia.htm

35. Huxtable CHA (1990) Ecological and embryological

studies of Microlaena stipoides (Labill.) R.Br. BSc(Hons) dissertation, University of New England, Armidale, NSW.

36. Institute for Aboriginal Development (1988) 'Punu:

Yankunytjatjara plant use: traditional methods for preparing foods, medicines, utensils and weapons from native plants.' (Angus and Robertson Publishers: Sydney). Scope: The region, people, language and plant use of the Yankunytjatjara. Perennial grass species: Eragrostis eriopoda, Panicum decompositum, Amphipogon caricinus Detail: Eragrostis eriopoda – tussock-forming grass, to 60 cm high, coarse hairy roots, dense woolly butt, found mainly on Spinifex sandplains and sandhills, and in mulga woodland. Also on river floodplains and levees, and on limestone rises. Seed ripening in autumn or winter after summer rains, ground to flour. Panicum decompositum – stout, tussock-forming, to 1 m high, most common on banks and floodouts of watercourses, seed ripening after summer rains, ground to a paste. Amphipogon caricinus – tufted grass, to 60 cm high, wiry stems, usually on sandy soils in association with Spinifex, not food.

37. Isaacs J (1987) 'Bush Food. Aboriginal food and

herbal medicine.' (Weldons: Sydney). Scope: Use of seeds of native grasses, shrubs, herbs and trees forming an important part of the Aboriginal diet. Perennial grass species: Astrebla pectinata, Brachiaria spp., Brachiaria miliiformis, Eragrositis eriopoda, Panicum decompositum Detail: Food grasses occur in Arnhem Land, Cape York and coastal regions, Lake Mungo in western NSW, Kimberly region, grasses of the Panicum, Brachiaria and Panicum genera seem to predominate and are common throughout the centre (particularly along watercourses, and in floodplains or mulga areas), Astrebla pectinata is found along the Darling River and used by the Bagundji people, grass seed is generally tiny: the largest, arm grass millet seed, is no more than 2 mm in diameter and most are smaller. Grass seed is ripe for harvesting from January to March. Most seeds are ground before eating. Panicum species – delicious seed for bread.

38. James KW (1985) Nutrients in native plant seeds —

2. In 'The food potential of seeds from Australian native plants. Proceedings of a colloquium held at Deakin University on 7 March 1984.' (Ed. GP Jones) pp. 46–57. (Deakin University Press: Victoria)

39. Johnson SB (1993) Seed germination and seedling

establishment of Microlaena stipoides (Labill.)

R.Br. BSc(Hons) dissertation, University of New England, Armidale, NSW.

40. Johnston WH, Mitchell ML, Koen TB, Mulham

WE, Waterhouse DB (2001) LIGULE: An evaluation of indigenous perennial grasses for dryland salinity management in south-eastern Australia. 1. A base germplasm collection. Australian Journal of Agricultural Research 52, 343–350.

41. Jones GP (1985) Can indigenous Australian plants be

seriously considered as potential food crops? In 'The food potential of seeds from Australian native plants Proceedings of a colloquium held at Deakin University on 7 March 1984'. (Ed. GP Jones) pp. 202–211. (Deakin University Press: Victoria). Scope: Summary of discussion at the above colloquium. Perennial grass species: none mentioned specifically. Detail: the need for an integrated research effort; the need for financial support of ecological research; what research is being conducted into indigenous plant foods; the uncertainty of research funding; co-ordinated effort is required perhaps using existing agencies; action to be taken; broad-acre farming of native plants?; nutritional advantage of native plants.

42. Kortt J (1985) Characteristics of the proteinase

inhibitors of Acacia seeds. In 'The food potential of seeds from Australian native plants. Proceedings of a colloquium held at Deakin University on 7 March 1984.' (Ed. GP Jones) pp. 120–146. (Deakin University Press: Victoria)

43. Lazarides M, Hacker J, Andrew M (1991)

Taxonomy, cytology and ecology of indigenous Australian sorghums (Sorghum Moench: Andropogoneae: Poaceae). Australian Systematic Botany 4, 591–635.

44. Loch DS, Harvey GL, Cole JP (1994) Seed

production of Themeda triandra (Kangaroo grass) — preliminary studies. In ‘National Workshop on Native Seed Biology for Revegetation — 24–26 August 1994 Perth WA.’ (Eds SM Bellairs, LC Bell) pp. 118–122. (Australian Centre for Minesite Rehabilitation Research/ The Chamber of Mines and Energy of Western Australia Inc.: Perth)

45. Lodder MS (1989) Biology and landscape potential

of Danthonia species. Master of Letters Thesis, University of New England.

46. Lodder MS, Groves RH, Müller WJ (1994) Early

seedling growth of three species of Danthonia as affected by depth of sowing and nutrient supply. Australian Journal of Botany 42, 543–554.

47. Lodge GM (1995) Native grass improvement by

selection. In 'Perennial Grasses: Proceedings of the second Australian perennial grass workshop 17–19 October Launceston Tasmania' pp. 50–64). Perennial grass species: Astrebla spp., Danthonia spp., Microlaena stipoides, Themeda triandra

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48. Lodge GM, Schipp AJ (1993) Effects of depth and

time of sowing on emergence of Danthonia richardsonii Cashmore and Danthonia linkii Kunth. Australian Journal of Agricultural Research 44, 1311–1322.

49. Low T (1988) 'Wild food plants of Australia.'

(HarperCollinsPublishers: Sydney). Scope: notes on various native plants used for food. Perennial grass species: Eragrostis eriopoda, Panicum australiense, Panicum decompositum, Panicum effusum Detail: Panicum decompositum – large, often pale, blue-green leaves, seeds are about 1.5 mm long, widespread outback grass found on low-lying ground in most habitats, especially common on heavy clay soils, seeds produced in late summer and autumn, ground to flour. Panicum australiense (annual?)– bunch panic, smaller plant, about 15 cm tall, often with reddish leaves, seed heads hidden among foliage. Panicum effusum – hairy panic, 20–60 cm tall, similar looking to P. decompositum. Eragrostis eriopoda – coarse wiry grass with rigid pointed leaves, readily identified at ground level by its white woolly butts, grows in clumps throughout the outback on red sandy soils, often as the dominant grass in mulga woodland, seeds as small as salt grains.

50. Lynch D (2000) (Ed.) 'West Australian Native Grass

Society Newsletter.' Vol. 1. 51. Magcale-Macandog DB, Whalley RDB (1991)

Distribution of Microlaena stipoides and its association with introduced perennial grasses in a permanent pasture on the northern tablelands of New South Wales. Australian Journal of Botany 39, 295–303.

52. Magcale-Macandog DB, Whalley RDB (1994)

Factors affecting the distribution and abundance of Microlaena stipoides (Labill.) R.Br. on the northern tablelands of New South Wales. Rangeland Journal 16, 26–38.

53. Maggiore PMA (1985) Utilisation of some

Australian seeds in edible food products. In 'The food potential of seeds from Australian native plants. Proceedings of a colloquium held at Deakin University on 7 March 1984.' (Ed. GP Jones) pp. 59–74. (Deakin University Press: Victoria)

54. Maiden JH (1975) 'The useful native plants of

Australia.' (Compendium: Melbourne, reprint of 1914 publication). Scope: indigenous vegetable food resources Perennial grass species: Panicum colonum, Panicum decompositum, Panicum trachyrachis, Microlaena stipoides

Detail: lists about 150 different species used for human food. Panicum colonum – erect stems for 2–8 feet high, very succulent, seeds used for bread, not endemic to Australia. Found in North Qld. Panicum decompositum – semi-aquatic species, tall, coarse and succulent, produces an abundance of feed, good forage plant, seeds in December and January, the grains pounded yield excellent food, small seeds, not endemic to Australia. All states except Tasmania – valuable in the Darling Downs (Qld). Panicum trachyrachis – valuable open pasture grass, quick growing, high biomass in summer, free seeder, seeds sometimes used by natives for food. Found in NSW, Qld and northern Australia. Microlaena stipoides – green all year, will live on poor soil, does not always freely seed. All states.

55. Malcolm B, Mitchell M, Crosthwaite J (1999)

Assessing the technical and economic potential of Microlaena stipoides in farming systems in southern Australia. SPIRT grant application.

56. Mitchell M, Rich E (1995) Native grasses realise their

potential. In 'Use it or lose it — 36th Annual Conference' pp. 187–188. (Grassland Society of Victoria: Victoria). Scope: potential of native grass species for agricultural and conservation use. Perennial grass species: Microlaena stipoides, Themeda triandra Detail: Summary of LIGULE project (Low Input Grasses Useful in Limiting Environments) 33 species (800 lines) were originally collected from 200 sites throughout Vic and NSW. Twenty species were assessed, including lines of Microlaena stipoides at five sites in Victoria and NSW on typical upland recharge zones, rainfall 500–700 mm, relatively acid soils. Assessed monthly for performance indicators such as survival, growth, flowering and seed production. No data in article.

57. Mitchell ML, Koen TB, Johnston WH, Waterhouse

DB (2001) LIGULE: An evaluation of indigenous perennial grasses for dryland salinity management in south-eastern Australia. 1. Field performance and the selection of promising ecotypes. Australian Journal of Agricultural Research 52, 351–365.

58. Munnich DJ, Simpson PC, Nicol HI (1991) A

survey of native grasses in the Goulburn District and factors influencing their abundance. Rangeland Journal 13, 118–129.

59. Murphy MA (1995) Vacuum harvesting of the native

grass Microlaena stipoides. In 'Proceedings of the 10th Annual Conference of the New South Wales Grassland Society'. Armidale, NSW p. 79. Scope: trialled two commercially available vacuum machines for harvesting Microlaena stipoides. Perennial grass species: Microlaena stipoides Detail:

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Harvesting techniques: 1. McCulloch Super Air Stream IV gas blower/vac which draws air through a centrifugal fan (McCulloch). 2. Hand harvested by shaking culms into a bucket (Hand 1). 3. Flymo electric vacuum which employs a venturi-vacuum system bypassing the fan (Flymo). 4. Hand harvested by shaking culms into a bucket (Hand 2).

One kilogram of Microlaena collected and harvested with each system produced the following weight of germinable spikelets: McCulloch 438 g Hand 1 744 g Flymo 950 g Hand 2 938 g

60. Murphy MA (1996) Biology and potential forage

value of Elymus scaber (R.Br.) A. Love. BSc(Hons) dissertation, University of New England, Armidale, NSW.

61. Murphy MA, Jones CE (1999) Observations on the

genus Elymus (Poaceae: Triticeae) in Australia. Australian Systematic Botany 12, 593–604. Scope: morphological characters of 19 specimens of Elymus, taxonomy, reproductive biology, geographic distribution, genome constitution. Perennial grass species: Elymus scaber, Elymus multiflora, Elymus rectisetus

62. Nolan M (1994) Harvest, germination and

establishment techniques for the revegetation of Themeda australis. Master of Horticulture, University of Western Sydney, Hawkesbury.

63. O'Connell J, Latz P, Barnett P (1983) Traditional and

modern plant use among the Alyawara of central Australia. Economic Botany 37, 80–109. Scope: descriptive summary of information on traditional and modern uses of native plants by Alyawara-speaking Australian Aborigines – including data on 157 species, 92 of which are used for food (36 for seeds). Descriptions of food plants cover form and distribution, collecting and processing techniques, caloric yields and dietary importance. Perennial grass species: Eragrositis dielsii, Eragrostis eriopoda, Panicum decompositum, Panicum effusum Detail: Alyawara territory is centred on the upper and middle Sandover River drainage about 240 km NE of Alice Springs. Seeds of grasses are all very small. The largest, those of armgrass millet (Brachiaria miliiformis) are no more than 2 mm in diameter and 3.2 mg in weight, producing about 1000 – 2000 seeds per plant (~ 2.3 g seed/plant). Detailed harvesting techniques. Eragrositis dielsii – short-lived perennial, not common, found on alluvial soils, especially stream banks, floodbanks. Seeds eaten. Available warmer months.

Eragrostis eriopoda – extremely common and widespread, often locally dominant. Seeds eaten. Available almost any season with sufficient rainfall. Panicum decompositum – common on alluvial soils, especially floodplains. Seeds eaten. Available warmer months. Panicum effusum – disturbed areas in mulga woodland. Seeds eaten. Available mainly summer.

64. Quinn JA, Hodgkinson KC (1983) Population

variability in Danthonia caespitosa (Gramineae) in responses to increasing density under three temperature regimes. American Journal of Botany 70, 1425–1431.

65. Quinn JA, Hodgkinson KC (1984) Plasticity and

population differences in reproductive characters and resource allocation in Danthonia caespitosa (Gramineae). Bulletin of the Torrey Botanical Club III (1), 19–27.

66. Robinson JB, Munnich DJ, Simpson PC, Orchard

PW (1993) Pasture associations and their relation to environment and agronomy on the Goulburn district. Australian Journal of Botany 41, 627–636.

67. Scott AW, Whalley RDB (1984) The influence of

intensive sheep grazing on genotypic differentiation in Danthonia linkii, D. richardsonii and D. racemosa on the New England Tablelands. Australian Journal of Ecology 9, 419–429.

68. Semple W, Koen T, Cole I (1999) Establishing native

grasses in degraded pastures of central western New South Wales. The Rangeland Journal 21, 153–168. Perennial grass species: native grasses in general Detail: native grasses are generally characterised by combinations of poor seed retention, chaffy seed structures, various types of dormancy and difficulty in obtaining uniform, good quality seed. Microlaena stipoides germination percentage (floret – one year old) was 87.5%, dormancy not noted. In field trial seed was sown at ~200 seeds/m row.

69. Sindel BM, Davidson SJ, Kilby MJ, Groves RH

(1993) Germination and establishment of Themeda triandra (Kangaroo grass) as affected by soil and seed characteristics. Australian Journal of Botany 41, 105–117.

70. Sindel BM, Groves RH (1991) Seed production

potential and domestication of Kangaroo grass (Themeda triandra). In ‘Native Grass Workshop Proceedings’. (Eds PM Dowling, DL Garden) pp. 171–172. (Australian Wool Corporation: Dubbo, NSW)

71. Smith SR, Jr., Whalley R (2002) A model for

expanded use of native grasses. Native Plants Journal 3, 38–49. Scope: a proposed model to be used as a guide for anyone interested in expanded use of grasses. Perennial grass species: native grasses in general

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Detail: a series of 12 steps starting with determining the need and choosing the appropriate species, progressing sequentially through collection strategies, potential breeding methods, seed production, developing an extension management package and developing markets. Each step is described in detail and relevant examples are given from North America and Australia.

1. Determine the need.

2. Select an appropriate species that meets this need.

3. Determine the breeding system and ploidy level of the selected grass species — knowledge of the breeding system of a grass species is critical before initiating a breeding or selection program.

4. Determine the geographic and ecological range of the species and assess the proportion of this range which needs to be sampled to meet the need – it is reasonable to assume that a widespread species would usually be a better prospect to fill a specific need than a species with a narrow range. When a native grass is being considered for expanded use, its range should first be surveyed through existing herbarium collections, species distribution maps in the literature and by surveying botanists, taxonomists and grassland managers.

5. Make a collection of the species over the selected geographic and ecological range – after determining geographic and ecological range, a collection strategy should be designed to collect plants and/or seed across this range, depending on the need and the proposed method for expanded use. Latitudinal ranges are important in terms of winter survival and day length requirements for flowering. The ideal collection of a desired species should include plant material from each of the distinct ecological environments across the selected range. If the full ecological range of a species is to be sampled then the final collection should also include sites with widely differing management histories in terms of grazing, fertiliser history, time since cultivation and fire management. Additional important features are position in the landscape, lithology and soil types.

6. Assess the genetic and morphological diversity of the species collection – diversity should be determined by characterising plant morphology from each collection site as individual genotypes and as composites of genotypes. Individual plant measurements include plant height and width, leaf characteristics, tiller characteristics, flowering date and components of seed yield. Morphological characterisation provides extensive information on each genotype and indicates the genetic diversity within and between collection sites. The use of molecular techniques allows assessment of diversity not subject to environmental effects.

7. Determine the limiting factors for expanded use of this species – must first identify the major limitations within a species before solutions can be developed that will allow expanded use. Overcoming limitations will range from breeding for specific traits to improved agronomic practices during establishment

and seed production. A major limitation for expanded use of native grasses is seed production. Most can be best described as conservative seed producers, partitioning photosynthate to maximise plant survival rather than seed production. The traditional method for overcoming limitations in seed yield is through breeding.

8. Formulate and implement an appropriate selection strategy for development and commercial release (eg. variety, cultivar, ecotype, ecovar, germplasm etc) – the following breeding techniques are all possibilities: mass selection (MS), recurrent phenotypic selection (RPS)(most common technique for cross-pollinated grass breeding), restricted recurrent phenotypic selection (RRPS), half-sib progeny selection (HS Progeny), clonal selection (vegetatively propagated material), superior genotype selection (self-pollinated and apomictic), a combination of these techniques, or other techniques. Australian selection programs with Microlaena stipoides used a modified form of RPS, whereby the superior genotype was selected over a series of generations. Single genotype selection was possible since self-pollination predominated (Clifford 1962).

9. Determine the system for registration and release to end users

10. Develop procedures for the successful establishment of seed production and commercial stands

11. Develop and produce an extension management package for seed growers and commercial end users

12. Develop national markets (international markets if appropriate)

72. Turner F (1891) The grasses of New South Wales:

Microlaena stipoides, R. Br. "Meadow rice grass". Agricultural Gazette of NSW 2, 22–23.

73. Vogel W (1995) The Hidden Asset: Native Pasture

Management. In 'Use it or lose it — 36th Annual Conference' pp. 188–189. (Grassland Society of Victoria: Victoria). Perennial grass species: Danthonia spp., Microlaena stipoides

74. Whalley RDB, Brown RW (1973) A method for the

collection and transport of native grasses from the field to the glasshouse. Journal of Range Management 26, 376–377.

75. Whalley RDB, Jones CE (1997) 'Commercialising the

Australian native grass M. stipoides.' Rural Industries Research and Development Corporation, RIRDC Publication No 97/34, Canberra. Also published as Whalley RDB, Jones CE (1998) 'Microlaena: Commercialisation and development of an agronomic package for Microlaena stipoides.' Meat and Livestock Australia., UNE.039. Scope: A report on commercialising the grass for turf, amenity and forage Perennial grass species: Microlaena stipoides

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Detail: native yearlong green perennial, indeterminate flowering species, found throughout the higher rainfall zones from the mountains of the Cape York Peninsula through New South Wales and Victoria to Tasmania, as well as in the wetter coastal districts of South Australia and the south west of Western Australia. Drought tolerant, more abundant on acid soils. Seed production occurs from December through to April. Seed yields up to 2.2 t/ha on small plots, 0.7 t/ha in larger blocks. Seeds usually weigh between 4 and 6 mg. Report looks at the selection of 3 varieties, efforts to increase the seed, patterns of seed production and seed harvesting technology. It also describes the sowing (depth, time), turfgrass management and forage trials, along with herbicide response.

76. Willemse LPM (1982) A discussion of the Ehrharteae

(Gramineae) with special reference to the Malesian taxa formerly included in Microlaena. Blumea 28, 181–194. Scope: Description and notes on the taxa (below) Perennial grass species: Ehrharta, Microlaena, Tetrarrhena, Petriella spp.

77. Williams OB, Lazarides M (1985) Australia. In 'Plant resources of arid and semi-arid lands'. (Eds JR Goodin, DK Northington) pp. 35–67. (Academic Press: Orlando). Scope: Plant resources of Australia’s arid interior Perennial grass species: Brachiaria spp., Dactyloctenium radulans, Eragrostis spp., Panicum spp. Detail: Seeds require considerable effort to collect and prepare but nevertheless are an important food item. Dactyloctenium radulans – button grass.

78. Winder P (1997) The agricultural potential of

Acacia species as a food source. http://www.agfor.unimelb.edu.au/STUDENTS/63875/paper.html

79. Woodland PS (1964) The floral morphology and

embryology of Themeda australis (R.Br.) Stapf. Australian Journal of Botany 12, 157–172.

80. Worsley P (1961) The utilization of natural food

resources by an Australian Aboriginal tribe. Acta Ethnographica Academiae Scientiarum Hungaricae 10, 153–190.

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B. Exotic perennial grasses with potential for use as a human food 1. Anon (1998) Perennial cereal rye: may beat barley.

Lethbridge Research Center Advance Summer 2. Anon (1999) Selected North Dakota and Minnesota

Range plants. http://www.greatplains.org/resource/1999/ndmnrngp/agrointe.html

3. Almouslem AB, Amleh N (1999) An intergeneric

hybrid between durum wheat and diploid wheatgrass Lophopyrum elongatum (Host) A. Love. Kuwait Journal of Science and Engineering 26, 143–155.

4. Becker R, Wagoner P, Hanners GD, Saunders RM

(1991) Compositional, nutritional and functional evaluation of intermediate wheatgrass (Thinopyrum intermedium). Journal of Food Processing and Preservation 15, 63–77.

5. Board of Science and Technology for International

Development (1996) Wild Grains. In 'Lost Crops of Africa Volume 1 Grains' pp. 251–272. (National Academy Press: Washington DC). Scope: Use of a vast number of wild grasses by African people for food, both historically and now. Consideration is given to developing wild grains as modern foods Perennial grass species: Aristida pungens (aka Stipagrostis pungens), Echinochloa colonum, Echinochloa pyramidalis (perennial?), Echinochloa stagnina, Eragrostis gangetica?, Panicum staginum, Panicum subalbidum (perennial?), Panicum turgidum, Setaria sphacelata, Sporobolus fimbriatus?, Stenotaphrum dimidiatum, Themeda triandra, Urochloa brizantha, Urochloa mosambicensis? Detail: Aristida pungens – deep roots, long leaves, black grains, extremely drought resistant Panicum turgidum – deep roots, clump-forming, loose tussocks 1 m or so in diameter, spreads by long looping stolons building up mats of vegetation, bears its seeds on panicles that rise above the mat, extremely drought resistant (30–250 mm rainfall) Panicum stagninum – does not produce a useful grain, yields a thick syrup used in confections Echinochloa stagnina – grows in floodplains to 3 m, historically seed has been harvested for food, mainly used for fodder when water level drops Setaria sphacelata – robust, usually tufted grass, seeds eaten as famine food

6. Cane S (1989) Australian Aboriginal seed grinding

and its archaeological record: a case study from the Western desert. In 'Foraging and Farming: the evolution of plant exploitation'. (Eds D Harris, G Hillman) pp. 99–119. (Unwin Hyman: London).

7. Christiansen J, Jorgensen JR, Jornsgard B, Stolen O

(1997) Regrowth in barley (Hordeum vulgare L.) and rye (Secale cereale L.). Acta Agriculturae Scandinavica Section B — Soil & Plant Science 47, 215–220.

8. Eubanks M (1993) A cross between Tripsacum

dactyloides and Zea diploperennis. Maize Genetics Cooperation Newsletter 39.

9. Gustafson J, Lukaszewski A (1985) Early seed

development in the annual and perennial Secale taxa. Canadian Journal of Genetics and Cytology 27, 134–142.

10. Harlan J (1989) Wild-grass seed harvesting in the

Sahara and sub-Sahara of Africa. In 'Foraging and farming: the evolution of plant exploitation'. (Ed. DRH Harris) pp. 79–98. (Unwin Hyman: London, UK). Scope: Covers a number of species of grasses harvested for their grains in Africa Perennial grass species: Aristida pungens, Panicum turgidum Detail: Aristida pungens – relatively tall (to 1.5 m), tufted, plume-like inflorescence, black grains, usually seeds are ground into flour. Panicum turgidum – deep-rooted, drought resistant, spreads by long slender stolons forming a loose mat that is very useful for erosion control, grains are rather large for a wild grass, seeds not suitable for bread.

11. Jackson L, Dewald C (1994) Predicting evolutionary

consequences of greater reproductive effort in Tripsacum dactyloides, a perennial grass. Ecology 75, 627–641.

12. Jackson W (1999) Developing high seed yielding

perennial polycultures as a mimic of mid-grass prairie. In 'Agriculture as a Mimic of Natural Ecosystems.' (Eds EC Lefroy, RJ Hobbs, MH O'Connor, JS Pate) pp. 1–55. (Kluwer: Dordrecht)

13. Jauhar Prem P, Chibbar Ravindra N (1999)

Chromosome-mediated and direct gene transfers in wheat. Genome 42, 570–583.

14. Jensen K (2002) ARS germplasm holdings of

grasses and legumes. http://www.ars.usda.gov/is/np/systematics/grass&legume.html

15. Jensen KB, Zhang YF, Dewey DR (1990) Mode of Pollination of Perennial Species of the Triticeae in Relation to Genomically Defined Genera. Canadian Journal of Plant Science 70, 215–226.

16. McDonald R, Stephen R (1979) Effect of sowing

and harvesting dates on dry matter production of autumn-sown Tama ryegrass, ryecorn and oats. New Zealand Journal of Experimental Agriculture 7, 271–275.

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17. Parfeniuk A (1981) Plant breeding in the Don area [Activities of the Don Breeding Center, cereals, grain legumes, perennial grasses and legume forages]. Stepnye Prostory. [Saratov : Ministerstvo sel'skogo khoziaistvo RSFSR] Sept 9, 28–31.

18. Piper J, Kulakow P (1994) Seed yield and biomass

allocation in Sorghum bicolor and F1 and backcross generations of S. bicolor x S. halepense hybrids. Canadian Journal of Botany 72, 468–474.

19. Piper JK (1998) Growth and seed yield of three

perennial grains within monocultures and mixed stands. Agriculture Ecosystems & Environment 68, 1–11. Scope: Growth and seed yield of 3 perennial species (2 grass, 1 legume) in monoculture, biculture and triculture treatments, on 2 soils differing in initial fertility, and over 5 years. Perennial grass species: Leymus racemosus (mammoth wildrye), Tripsacum dactyloides (eastern gamagrass) Detail: Leymus racemosus – rhizamotous C3 species native to Bulgaria, Romania, Turkey, former Soviet Union. Its grain has been gathered by Asian and European people, especially in drought years when annual grain crops have failed. Reproductive tillers grow to about 1.5 m high and maximum seed yields have ranged from 51 to 83 g/m2. Tripsacum dactyloides – large C4 bunchgrass native from the southeatern United States and Great Plains southward to Bolivia and Paraguay. Canopy height ranges from 1 to 2 m, reproductive tillers can exceed 2 m high. Gamagrass is an excellent forage species, it also shows much promise as a grain crop for human consumption. Grain contains 27–30% protein, 7% fat and has baking properties similar to those of maize. Seed yield is low 40–100 g/m2.

20. Reimann-Philipp R (1986) Perennial spring rye as a

crop alternative. Journal of Agronomy and Crop Science 157, 281–285.

21. Reimann-Philipp R, Gordon-Werner E (1984)

Investigation of cytological tests for improving the fertility of a tetraploid perennial spring rye (S. cereale x S. montanum). Zeitschrift fur Pflanzenzuchtung 92, 198–207.

22. Vogel KP, Shearman RC (1996) Perennial grasses:

new applications and uses. In 'Progress in new crops.' (Ed. J Janick) pp. 263–270. (ASHS Press: Virginia)

23. von Bothmer R, Jacobsen N, Jorgensen R, Linde-

Laursen I (1984) Haploid barley from the intergeneric cross Hordeum vulgare x Psathyrostachys fragilis. Euphytica 33, 363–367.

24. Wagoner P (1990) Perennial grain development: past

efforts and potential for the future. Critical Reviews in Plant Sciences 9, 381–408. Scope: Efforts to develop perennial grain crops, initiated during the 20th century, are discussed —

including perennialising annual grains and domesticating perennial grass species. The advantages and disadvantages of these two approaches to perennial grain development are discussed, with examples. Perennial grass species: Thinopyrum intermedium (intermediate wheatgrass), Tripsacum dactyloides (eastern gamagrass), Distichlis palmeri (saltgrass), Leymus spp. (wildrye) Detail: excellent review covering: • The benefits of perennial grain development • Historical perspectives • The development of perennial grain crops – factors to consider • Development of economically viable perennial grains • Approaches to the development of perennial grains • Efforts to perennialise annual grains (wheat, rye, maize, sorghum) • Efforts to domesticate perennial grasses (intermediate wheatgrass, wildrye, saltgrass, eastern gamagrass)

25. Wagoner P (1990) Perennial grain: A new use for

intermediate wheatgrass. Journal of Soil and Water Conservation 45, 81–82.

26. Wagoner P (1994) Perennial Grain: a new use for

intermediate wheatgrass. New Crop News 4, 3–4. Scope: discusses potential of intermediate wheatgrass as a commercially viable perennial grain crop: grain production, economics, cultivar development, marketing, supply & demand, prospects for the future. Perennial grass species: Thinopyrum intermedium (intermediate wheatgrass) Detail: Intermediate wheatgrass or Wild Triga– perennial relative of wheat, high protein grain (20%), mild sweet nutty flavour, gluten free, widely adapted throughout US and Canada to areas with annual rainfall of 375 mm or more, currently used as a forage and for erosion control along roadways and mine reclamation sites. Grain yields at the Rodale Institute in the first year range from 2.75–3.85 t/ha, which declines in subsequent years.

27. Wagoner P (1995) Wild Triga. Intermediate

wheatgrass. New Crop FactSHEET. 28. Wagoner P, Schauer A (1990) Intermediate

wheatgrass as a perennial grain crop. In 'Advances in new crops: proceedings of the first national symposium 'New crops: research, development, economics', Indianapolis, Indiana, USA, 23–26 October 1988'. (Eds J Janick, JE Simon) pp. 143–145. (Timber Press: Portland, Oregon, USA). Scope: The development of perennial grains such as intermediate wheatgrass for environmental benefits through the reduction of soil erosion and a reduction in the inputs necessary to produce annual crops. Perennial grass species: Thinopyrum intermedium (intermediate wheatgrass)

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Detail: long term research project commenced in 1983 to investigate the potential of developing a perennial grain cropping system. Close to 100 species of perennial grasses were evaluated to identify those with good potential for development into perennial grain crops using 10 selection criteria (outlined in paper). Intermediate wheatgrass, a perennial relative of wheat, was selected as having good potential. 250 accessions were acquired and tested in small plots to determine seed production capabilities. Seed size of 150 accessions ranged from 3.6–7.2 g/1000 seeds. Other larger scale plots varying planting date and seeding rates have been tested. Also nutritional and food use evaluations.

29. Wang R (1986) Diploid perennial intergeneric

hybrids in the tribe Triticeae. I. Agropyron cristatum

x Pseudoroegneria libanotica and Critesion violaceum x Psathyrostachys juncea. Crop Science 26, 75–78.

30. Wang R (1988) Coenocytism, ameiosis, and chromosome diminution in intergeneric hybrids in the perennial Triticeae. Genome 30, 766–775.

31. Weibull Jens HW (1993) Bird cherry-oat aphid

(Homoptera: Aphididae) performance on annual and perennial temperate-region grasses. Environmental Entomology 22, 149–153.

32. Wilson D, Breese E, Valentine J (1988) Selection

strategies for quality breeding. Hodowla Roslin Aklimatyzacja i Nasiennictwo 32, 1–2.

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C. Further detail on Microlaena stipoides 1. Aldous D, Chivers I (1996) Agronomic and quality factors of weeping grass (Microlaena stipoides). New Zealand Turf

Management Journal February, 31–32. • Microlaena stipoides grows naturally throughout the higher rainfall coastal and tableland regions of eastern

Australia, as well as coastal south and western Australia. In New Zealand it grows throughout the North Island and is a dominant native species on many of Auckland’s volcanic cones.

• slightly rhizomatous and stoloniferous perennial grass, growing up to 15 cm tall (seed heads up to 40 cm) with a fibrous root system.

• frost and drought tolerant. • persists in heavy shade. • greenhouse experiment with a number of accessions. Measurements taken: tiller number/pot, tiller numbers

emerging from rhizome growth/pot, leaf length/width, leaf width, culm length. 2. Anon. (1995a) Variety: 'Griffin' syn. '703.6.12'. Application no. 95/052. Plant Varieties Journal 8, 22. 3. Anon. (1995b) Variety: 'Shannon' syn. '17.2.6.5.12'. Application no. 94/124. Plant Varieties Journal 8, 22. 4. Anon. (1995c) Variety: 'Wakefield' syn. '39.1.8.2.5'. Application no. 94/125. Plant Varieties Journal 8, 22.

Shannon Wakefield Griffin

Growth Habit (1=prostrate, 5 = erect) 3 3 2

Plant Height (cm) 19.1 ± 5.2 27.6 ± 4.3 15.3 ± 5.8

Leaf Colour Light green Medium green Medium green

Leaf attitude (1=upright, 5 = drooping) 3.8 2.7 2.2

Flag leaf width (mm) 4.08 ± 0.79 3.85 ± 0.69 3.19 ± 0.80

Flag leaf length (mm) 51.2 ± 12.3 56.7 ± 11.9 33.3 ± 10.0

Inflorescence length (mm) 119.2 ± 17.1 160.6 ± 24.9 140.2 ± 24.9

Number of spikelets/inflorescence 28.3 ± 5.2 46.4 ± 9.4 25.7 ± 6.3 5. Archer K, Robinson G (1988) Agronomic potential of native grass species on the Northern Tablelands of New South

Wales. II. Nutritive value. Australian Journal of Agricultural Research 39, 425–436. • laboratory analyses for in vitro organic matter digestibility (OMD), nitrogen (N) and phosphorus (P). Sheep fed. • late spring – in vivo OMD 65.9% (Microlaena stipoides) cf. 71.8% (white clover) – Microlaena was significantly

higher than the other grasses. • summer – in vivo OMD 66.8% (Microlaena stipoides). • overall Microlaena had the highest N concentration of any of the grasses, and had especially high levels in spring

with up to 4.6 g N/100 g OM in the green leaves. • microlaena had high levels of P in unsorted and green leaf fractions, ranging from 0.38 g P/ 100g OM in the

green leaves in spring to 0.59g P/ 100g OM in the green leaves in autumn. • microlaena – high year round digestibility of green leaves, high N and P concentrations BUT has low growth

rates during winter and is limited in its distribution to more fertile, protected areas. 6. Baxter A (Ed.) (2000) 'Native grasses information kit.' (Agriculture Western Australia: Perth).

• Microlaena stipoides seed is produced in Nov/Dec and, in response to summer and autumn rains, from January to May.

• remains green throughout the year. • more commonly seen in damp or semi-shade areas. • major component of Mount Lofty Ranges woodlands. • 30–70 cm. • high frost and drought tolerance. • as many as 100 viable cuttings can be obtained from large plants by simple cutting the rhizomic root system at

each root node. • seed size looks to be 10 mm x 1.5 mm. • seed is obtained by stripping florets off the long narrow panicle at the end of the weeping seed stalk. As much

time as practicable should be allowed for the seed to mature before harvesting. When pinched between the thumbnail and forefinger viable seed will feel stiff and firm in comparison to the hollow and easily bent infertile florets.

• a good average crop should yield 10 g dry seed/m2 with 70–80% viability.

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7. Campbell CS, Quinn JA, Cheplick GP, Bell TJ (1983) Cleistogamy in grasses. Annual Review of Ecology & Systematics 14, 411–441. • cleistogamy (CL) – self-fertilisation in an enclosed flower – intensifies the potential advantages of inbreeding; it

may also be important in altering seed size, germinability, dispersibility, and susceptibility to fire and animal predation (Campbell 1982, Dobrenze and Beetle 1966, Wilvert 1982).

• CL is an evolutionary derived and complex syndrome of morphological, developmental, genetic and environmental components. The variability in the expression of CL in grasses may be considered from the perspective of any one component or a combination of them.

• the structures adapted for the protection of aerially exposed, chasmogamous (CH) spikelets, florets and fruits, and for fruit dispersal, are often either reduced in size and/or number or lost altogether.

• four types of CL – Microlaena stipoides is Type 1 – Sheath fertilisation: inflorescences or spikelets remain within the leaf sheathes of the middle or uppermost part of the stem for fertilisation and for fruit maturation, or they may be exserted from the sheath at some time after fertilisation.

• relatively meagre amounts of pollen have been reported in the CL anthers of Microlaena. The anther length is strongly correlated with the number of pollen grains in the anther (Clifford 1962). Since grass flowers have only one ovule, the anther number and length provide a good estimate of the pollen-ovule ratio and thereby reflect variation in breeding systems.

• CL spikelets either in eventually emergent panicles or solitary and remaining within sheaths. • awns: CH and emergent CL longer than those of spikelets remaining in sheaths. • paleas: CH more than 6 mm long, CL less than 5 mm. • lodicules: CH 2–3 mm, CL 0–small. • anthers: CH – usually 4 (occasionally 1, 2, 3 or 6); 2.4–4 mm. CL usually 2, sometimes 4 in CL spikelets

remaining in sheaths, and usually 4 in emergent CL spikelets, 0.7–1.4 mm. • stigmas: CH 2.2 mm, CL 0.9–2.3 mm.

8. Chivers IH, Aldous DE (1996) Response of weeping grass (Microlaena stipoides) to pre-emergence and post-

emergence herbicides. In 'Proceedings of the 3rd ATRI Turf Research Conference'. Brisbane, Qld & Sydney NSW pp. 80–87. • Trifolium repens (white clover) can be a serious weed of weeping grass. • studies were conducted with weeping grass to investigate the phytotoxicity caused by trifluralin (as Treflan),

dithiopyr (as Dimension), pendimethalin (as Stomp), bensulide (as Exporsan), 2,4-D as the sodium salt formulation (as Yates Lawn Weed & Feed) and MCPA and dicamba as the dimethylamine salt formulation (as Hortico Clover & Bindii).

• pre-emergence herbicide field screening trial and post-emergence herbicide pot trial. • a pre-emergence spring application of pendimethalin (2.0 and 4.0 l/ha) or an autumn application of trifluralin (1.5

l/ha) was effective as pre-emergent herbicides on the germinating weedy growth, with limited phytotoxic effects recorded on the emerging weeping grass seedlings. The other herbicides were less successful (phytotoxic to weeping grass).

• post-emergence study: a single application of 2,4-D applied at 0.5 x, or at the recommended rate, did not have a significant phytotoxic effect on the weeping grass or the white clover. Neither did a repeat application. MCPA and dicamba at 0.5 x and the recommended rate did have a significant phytotoxic effect on both weeping grass and white clover.

9. Cochrane D (1998) Native perennial grasses for Western Australia. Land Management Society Newsletter November,

11–12. • Donald Cochrane is a farmer from Duranillin in south western WA • identifies Microlaena stipoides as having a degree of agricultural or amenity benefit. • weeping grass has a wide geographic distribution throughout southern Australia having the ability to adapt to a

wide range of soils and rainfall. The fact that some of the best natural stands of weeping grass as seen within the drip line of eucalypts highlights its versatility.

• research in eastern Australia (Simpson, pers comm.) – protein 14.8%; yield 8.3t/ha/yr. • seed kernel of weeping grass is very similar to a small grain of rice. • strong colonies can still be found in paddocks in south west WA around trees and fence lines where spraying and

cultivation have not taken place. 10. Cole I, Waters C (1997) Harvesting and sowing native grasses. In ‘Proceedings of the Twelfth Annual Conference of

the Grassland Society of New South Wales’. pp. 95–103. • harvest yield using brush harvester 153.9 kg over 14 ha = 10.9 kg/ha • mean florets/inflorescence before (11) and after (3) a single pass with brush harvester (73% stripped).

11. Connor H, Matthews B (1977) Breeding systems in New Zealand grasses. 7. Cleistogamy in Microlaena. New Zealand

Journal of Botany 15, 531–534. • microlaena is a small genus of grasses occurring in New Zealand, Australia, New Guinea, Indonesia and Hawaii. • flowers in emergent panicles of Australian Microlaena stipoides are of two readily distinguishable kinds,

chasmogamous (CH) and cleistogamous (CL) (Clifford 1962). • in CH flowers the anthers and lodicules are much larger than those of CL flowers.

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• the inflorescence in microlaena is a panicle or raceme, with numerous pedicellate spikelets. The spikelet, subtended by two small persistent glumes, consists of two awned, sterile lemmata and one perfect floret.

• Microlaena stipoides, in addition to panicles of CH flowers, produces panicles with solely CL flowers where fertilisation and seed formation precede inflorescence emergence.

• CL flowers – two types – aerial spikelets, clandestine spikelets – size and shape are quite different. The awns of clandestine spikelets are 2–3 mm long relative to 19–20 mm in aerial spikelets. The clandestine spikelet is asymmetric, with a marked groove where the spikelet lies against the internode. (more detail in the paper).

• several floral features distinguish CH and CL flowers. In general, flowers in clandestine spikelets have fewer anthers than flowers in aerial spikelets. Anthers in aerial and clandestine CL flowers of Microlaena stipoides are the same length, and so too are stigma styles (which are about half as long as those of CH flowers).

• pollination of aerial and clandestine CL flowers – pollen grains germinate within the anther, and pollen tubes grow out through the anther wall and on to the stigmata.

Anthers/flower CH Aerial CL Clandestine CL

1 1 2

2 1 25 16

3 20 20

4 16 14

Mean anther length (mm) 2.49±0.08 0.3 ±0.01 0.34±0.01

Mean stigma-style length (mm) 2.39±0.12 1.3 ±0.05 0.96±0.05

Mean caryopsis length (mm) 4.52±0.12 3.58±0.17 3.74±0.34

Seed is produced by Microlaena stipoides in: • aerial CH flowers which, though probably self-fertile, may also be cross-fertilised. • aerial CL flowers where obligate self-fertilisation and seed development occur before inflorescence

emergence. • obligatory CL flowers in axillary, solitary, clandestine spikelets. These do not seem to be very frequent,

though they may occur at three consecutive nodes. • the co-occurrence in a single plant of Microlaena stipoides of aerial panicles of either CH or CL flowers and

of axillary, clandestine, CL spikelets appears unrelated to any ecological condition. 12. Connor HE (1979) Breeding systems in the grasses: a survey. New Zealand Journal of Botany 17, 547–574.

• grass flowers consist of three separate floral structures: the lodicules, succeeded by one or two (or even more) alternating staminal whorls, and the uniloculate terminal gynoecium. The floral organs are enclosed by an anthoecium – the lemma and palea; collectively the flower and the bracts are called a floret.

• lodicules – the main role is in opening the floret at anthesis. Commonly there are 2 lodicules in grasses and 3 in bamboos.

• in microlaena lodicules are absent or so small as to be almost unnoticed in CL flowers but larger in CH flowers (Clifford 1962).

• in most grass flowers there are 3 stamens, though stamen number varies. • CL clandestine spikelets occur at ground level. Usually they are very different from spikelets on aerial borne

inflorescences. Clandestine spikelets occur in Microlaena. • CL in aerially borne inflorescences may be facultative or obligate, though most would be best classified as

facultative because CH flowers are usually reported in the same species. “Predominantly cleistogamic” is used in some descriptions indicating the presence from time to time of CH flowers.

• evidence of environmental control of CL eg. Brown (1952) showed in Stipa leucotricha that CL could be a response to low soil moisture.

• mode of pollination: in Microlaena pollen germinates in the theca, grows out through the walls of the pollen sac and on to the stigmas (Clifford 1962, Connor 1977).

13. Earl JM, Whalley RDB, Jones CE (1994) Variation in the components of seed yield and germination requirements

among three accessions of Microlaena stipoides. In 'Eighth Biennial Rangeland Society Conference'. Katherine, Northern Territory pp. 179–180. (Australian Rangeland Society, Alice Springs). • seed production and components of seed yield of three accessions of Microlaena stipoides were measured over

one growing season. • final seed yield did not differ among accessions although there were differences in partitioning. • the accession with the greatest number of inflorescences had the least number of spikelets/inflorescence, while

the accession with the least inflorescences/plant had the most spikelets/inflorescence. • seed yields ranged from 1.7–2.2 t/ha.

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14. Groves RH, Whalley RDB (2002) Grass and grassland ecology in Australia. Flora of Australia 43, 157–182. • generally an inverse relationship between size of individual seeds and number of seeds produced per plant. • inherent in the dispersal mechanisms of many Australian grasses is the development of abscission layers (made

up of layers of cells that die as the seed matures), structures that ensures that seeds disperse as they ripen. In Microlaena stipoides an abscission layer develops below the individual dispersal unit which then falls from the inflorescence as it ripens. The spikelets fall as a unit with the abscission layer forming above the glumes where there is only one fertile floret per spikelet.

• many Australian grasses are perennials that flower indeterminately whereby seeds are formed and ripen over an extended period of time. This general feature is ecologically advantageous but poses problems if seed is to be harvested. Microlaena stipoides produces inflorescences throughout the summer, provided soil moisture is available. Rainfall patterns have a marked effect on flowering and if there is a wet spring followed by a dry summer, they may appear to show determinate flowering. In other years, depending on the rainfall events, such grasses may have one flowering period in the spring and another in the autumn with few flowers in between.

• total mass of herbage produced in the summer is much higher than in the winter period, hence ‘year-long green’. • ancillary structures of Microlaena seeds are difficult to remove without damage to the caryopses. Complete

cleaning of the dispersal unit to naked caryopses my reduce dormancy in some species (Lodge 1981). There is some evidence that seed longevity may be reduced when dispersal units are cleaned completely so that only naked caryopses remain (Grice 1995).

• dispersal units of Microlaena stipoides bear straight or slightly curved awns that are not hygroscopic. Peart (1984) showed that these awns have an aerodynamic function that ensures that the dispersal units land with the callus end down when shed from the parent plant. Peart (1984) showed further that the backward-facing hairs on the callus help to anchor the seed to the ground and that if such dispersal units remain in an upright position have a high probability of germinating and the seedlings becoming established. Results of more recent work indicates that water uptake through the callus end of the dispersal unit is faster than for the rest of the structure (M. Paterson, pers. comm.). Mechanical treatment to remove the awns without damaging the caryopses is very difficult.

• apomixis is the production of viable seeds by plants without the union of gametes ie. seeds without sex. An embryo sac is formed inside the ovary from an unreduced cell and the resultant embryo usually has a genotype identical to that of the parent plant. Pollination is usually necessary for the development of apomictic embryo sacs and is also necessary for the development of the endosperm (Asker and Jerling 1992).

15. Lodge GM (1995) Native grass improvement by selection. In 'Perennial Grasses: Proceedings of the second Australian

perennial grass workshop 17–19 October Launceston Tasmania' pp. 50–64). • history of native grass domestication in Australia, features of major genera being studied, selection procedures

and criteria used, and the current status of native grass improvement programs. • Microlaeana stipoides var. stipoides

• yearlong green perennial grass • somewhat frost tolerant • major growth period is during summer provided soil water is available (Robinson 1988) • good drought survival • tolerant of shade and is commonly found beneath tree canopies or in lightly timbered areas • tolerant of low soil pH and high levels of both aluminium and manganese (Mitchell et al. 1992) • resistant to defoliation • majority of inflorescences have cleistogamous, self-pollinated spikelets with no evidence of apomixis

(Clifford 1962) • following hot conditions in summer plants may produce a few chasmogamous spikelets (Huxtable 1990) • to date, all hexaploid (4n = 48) which behave as functional diploids with a basic chromosome number

x=12 16. Magcale-Macandog D, Whalley R (1991) Distribution of Microlaena stipoides and its association with introduced

perennial grasses in a permanent pasture on the Northern Tablelands of New South Wales. Australian Journal of Botany 39, 295–303. • Microlaena stipoides is one of the few Australian native grasses that provide forage during the critical winter-

early spring period on the Northern Tablelands of NSW (Robinson 1988). • year-long green perennial – remains green during winter and has an indeterminate growth pattern. • rapid growth and flowering whenever soil water is available from spring through autumn. • high grazing value (Lodge 1989). • protein content of its regrowth is generally higher in autumn, winter and spring than in the summer months. • the highest digestibility (63–79%) of the forage occurs during winter and spring and then declines in the late

summer and autumn months (from 45–67%) (Archer 1988). 17. Magcale-Macandog D, Whalley R (1994) Factors affecting the distribution and abundance of Microlaena stipoides

(Labill.) R.Br. on the Northern Tablelands of New South Wales. The Rangeland Journal 16, 26–38. • Microlaena stipoides is a perennial species which retains a significant proportion of green leaves year-round

(Robinson 1988), has high herbage production (Begg 1959) and thus provides forage during the winter-early spring period when feed is scarce (Robinson 1988).

36

• appears to be more common at higher elevations in the eastern parts of the Northern Tablelands (Lodge 1989). • if present in a paddock then it almost invariably occurs under trees, or if there are no trees, on the margins of

sheep camps (Whalley et al. 1978). • able to tolerate and survive on strongly acid soils (Munnich 1991). • adaptable to a wide range of soil fertility levels. It can be found on phosphate-fertilised and phosphate-poor soils. • very responsive to added nitrogen (Vieira 1980).

18. Maiden JH (1975) 'The useful native plants of Australia.' (Compendium: Melbourne).

• Microlaena stipoides – weeping grass, meadow rice grass, green all year, will live on poor soil, does not always freely seed. All states.

• an analysis in the spring resulted in: Albumen 1.66 Gluten 9.13 Starch 1.64 Gum 3.25 Sugar 5.05

19. Malcolm B, Mitchell M, Crosthwaite J (1999) Assessing the technical and economic potential of Microlaena stipoides

in farming systems in southern Australia. ARC Proposal — SPIRT Application • aim: to evaluate the technical and economic potential of the perennial native grass Microlaena stipoides in

farming systems in South Australia. • potential for grain production – produces a grain similar in size to rice. • as part of a whole farm/catchment approach, Microlaena stipoides could have a role in helping to reduce and

prevent problems of rising water table and salinity, erosion, structure decline and acidification. • native, year-long green perennial grass that grows in Australia’s high rainfall zone, from the mountains of the

Cape York Peninsula through New South Wales and Victoria to Tasmania, as well as in the wetter coastal districts of South Australia and Western Australia.

• currently there are four cultivars that have been registered: Griffin, Flinders, Wakefield and Shannon. None have been selected for their grain characteristics.

• technologies exist for the harvesting of grain from existing stands of Microlaena stipoides and grain cleaning technologies are also developed.

• tolerance to acid soils. 20. Mitchell M, Koen T, Johnston W, Waterhouse D (2001) LIGULE: an evaluation of indigenous perennial grasses for

dryland salinity management in south-eastern Australia. 2. Field performance and the selection of promising ecotypes. Australian Journal of Agricultural Research 52, 351–365. • initial evaluation of a large collection of Australian perennial native grasses – 807 accessions (37 species). • identify accessions that may be useful for pastoral purposes and for controlling land degradation on hill lands in

the high (>500 mm) rainfall zone of south eastern Australia. • Microlaena stipoides is a C3 species. • most active in summer, semi-erect/tussock, seed head density — dense, seed retention — shattering, reasonably

palatable. • seed shedding and inaccessible seed heads are recognised as barriers to commercial seed production in

Microlaena stipoides. • palatability may also be an issue in grazed pastures. • LIG183 and LIG704 showed good seed retention, dense inflorescences and an upright habit as well as good

winter growth and palatability. 21. Munnich DJ, Simpson PC, Nicol HI (1991) A survey of native grasses in the Goulburn District and factors influencing

their abundance. The Rangeland Journal 13, 118–129. • survey of natural and improved paddocks on 34 farms over winter within a 60 km radius of Goulbourn on the

Southern Tablelands of NSW. • Microlaena stipoides was relatively abundant (16%) on the natural paddocks. Cultivation did not affect its

abundance on the improved pastures. It was also more prevalent in uncleared than cleared paddocks. • the abundance of Microlaena stipoides was affected by pH, with percentage frequency increasing as pH declined

(down to pH 4.4 (CaCl2)). 22. Murphy MA (1995) Vacuum harvesting of the native grass Microlaena stipoides. In 'Proceedings of the 10th Annual

Conference of the New South Wales Grassland Society'. Armidale, NSW p. 79. • vacuum harvesting of the native grass Microlaena stipoides. • trialed two commercially available vacuum machines for harvesting Microlaena stipoides. • harvesting techniques:

a. McCulloch Super Air Stream IV gas blower/vac which draws air through a centrifugal fan (McCulloch). b. Hand harvested by shaking culms into a bucket (Hand 1). c. Flymo electric vacuum which employs a venturi-vacuum system bypassing the fan (Flymo). d. Hand harvested by shaking culms into a bucket (Hand 2).

37

• one kilogram of Microlaena collected and harvested with each system produced the following weight of germinable spikelets:

a. McCulloch 438 g b. Hand 1 744 g c. Flymo 950 g d. Hand 2 938 g

23. Robinson J, Munnich D, Simpson P, Orchard P (1993) Pasture associations and their relation to environment and

agronomy in the Goulburn district. Australian Journal of Botany 41, 627–636. • methodology as per Munnich (1991). • data was analysed using numerical classification methods to identify species associations particularly the

agronomic and environmental factors associated with the abundance of Danthonia spp. and Microlaena stipoides. M. stipoides was similarly correlated with sown grass species but had no correlation with annual grasses.

24. Robinson G, Archer K (1988) Agronomic potential of native grass species on the Northern Tablelands of New South

Wales. I. Growth and herbage production. Australian Journal of Agricultural Research 39, 415–423. • herbage mass and relative growth rate of six perennial native grasses compared with two introduced temperate

perennial grasses. • grasses sown at a density of 20 plants/m². • microlaena had a higher growth rate in late spring-summer but was lower during the winter months. Comparison

of the leaf and total green production shows the ability of Microlaena to produce a higher proportion of leaf during the spring.

25. Turner F (1891) The grasses of New South Wales: Microlaena stipoides. The Agricultural Gazette of New South Wales

2, 22–23. 26. Turner F (1921) 'Australian grasses and pasture plants : with notes on native fodder shrubs and trees.' (Whitcombe &

Tombs: Melbourne). • Microlaena stipoides (meadow rice grass) – found in all states from the coast to a considerable distance inland;

more plentiful on the coast. • strong root system enabling the grass to withstand long periods of dry weather. • yields the greatest bulk of herbage in rich meadow land. • under favourable conditions it sometimes attains a height of 3 feet, but generally it grows from 1–2 feet high. • may be easily recognised by its vivid green leaves, which it retains throughout the year in ordinary seasons. • superior pasture grass – its rich herbage is greatly relished by all herbivora; even under close grazing it will

maintain a dense turf. • if cut when in flower it makes an excellent hay. • seeds are freely produced and usually ripen during the summer and autumn. • albumen 1.66; gluten 9.13, starch, 1.64, gum 3.25, sugar 5.05% (Mueller and Rummel).

27. Whalley R, Brown R (1973) A method for the collection and transport of native grasses from the field to the

glasshouse. Journal of Range Management 26, 376–377. • a simple method for transplanting native grasses from the field to the glasshouse with negligible mortality:

1. entire plant is carefully removed from the soil with a shovel or similar tool, including a relatively undisturbed ball of soil around the roots.

2. entire plant is immediately placed in a large plastic bag and the leaves and culms clipped to 10% of their original height.

3. a small quantity of water is added to dampen (but not saturate) the soil, and the plastic bag is sealed and labelled.

4. plants are shaded as much as possible from direct solar radiation during transport. In the glasshouse the plants are placed in pots, adding a soil mix if necessary, and then watered.

• a number of different species have been successfully collected using this method (Microlaena not mentioned). These plants were collected successfully at different times during the growing season, including periods of active growth.

• better than direct potting in the field. • plants have been stored in plastic bags for several days in a vehicle without suffering severe mortality.

28. Whalley RDB, Jones CE (1997) 'Commercialising the Australian native grass M. stipoides.' Rural Industries Research

and Development Corporation, RIRDC Publication No 97/34, Canberra. Also published as Whalley RDB, Jones CE (1998) 'Microlaena: Commercialisation and development of an agronomic package for Microlaena stipoides.' Meat and Livestock Australia., UNE.039. • a report on commercialising the grass for turf, amenity and forage–looks at the selection of 3 varieties, efforts to

increase the seed, patterns of seed production and seed harvesting technology. It also describes the sowing (depth, time), turfgrass management and forage trials, along with herbicide response.

38

• found throughout the higher rainfall zones from the mountains of the Cape York Peninsula through New South Wales and Victoria to Tasmania, as well as in the wetter coastal districts of South Australia and the south west of Western Australia.

• native yearlong green perennial grass with a very flexible breeding system • drought tolerant, more abundant on acid soils. • indeterminate flowering species which continually produces inflorescences from early December through April • majority of inflorescences produced by Microlaena stipoides plants bear cleistogamous (CL) flowers with

pollination occurring within the closed lemma and palea (Clifford 1962) but there is no evidence for apomixis within these CL flowers (Huxtable 1990). Occasionally, inflorescences bearing chasmogamous (CH) flowers are produced with the potential for outcrossing (Clifford 1962). The result is that Microlaena stipoides behaves as a self-pollinated species provided seed is only collected from CL inflorescences and selection blocks can be satisfactorily grown in close proximity. The CH flowers usually appear for a short period in January and so care must be exercised to avoid collecting seed from these inflorescences if blocks of other varieties are producing CH flowers at the same time.

• seeds enclosed in ancillary structures including 2 scabrid awns. The term ‘seed’ is used to refer to the dispersal units, which comprise the actual seed plus the ancillary structures and awns.

• seeds usually weigh between 4 and 6 mg and should be sown at about 10 mm depth. • individual caryopses are ~6 mm long, weigh 2–4 mg and the embryos project about 0.7 mm beyond the end of

the endosperm. Each caryopsis is enclosed in a lemma and palea which are, in turn, enclosed in two awned sterile lemmas. The glumes are minute and an abscission layer forms above the glumes at maturity resulting in early shedding of the dispersal units.

• seeds gave a wide temperature range for germination between about 10°C to 35°C. The seedlings grow relatively slowly for some weeks.

• seed ripens progressively from the top of the inflorescence and tend to fall as they ripen. • 3 accessions – no significant difference in time to initial production, rate of seed production or maximum

production. The pattern of seed production varied. • seed yields up to 2.2 t/ha on small plots, 0.7 t/ha in larger blocks.

29. Willemse LPM (1982) A discussion of the Ehrharteae (Gramineae) with special reference to the Malesian taxa

formerly included in Microlaena. Blumea 28, 181–194. • Tribe: Ehrharteae; Genera: Ehrharta, Microlaena, Tetrarrhena and Petriella. • spikelets pedicelled, laterally compressed, 3-flowered • rachilla articulating above the glumes • lower two florets are sterile, epaleate • upper floret bisexual, paleate • Microlaena stipoides specifically: • glumes minute to short (usually <4 mm) • rachilla-process usually absent, occasionally present in some specimens of M. avenacea and M. giulianettii • sterile lemmas glabrous except for the bearded stipe, without basal appendages, not wrinkled, apex contracted

into a short to long false awn • palea 1-nerved • stamens usually 4 or 2 but 3, 1 or 6 have also been observed in M. stipoides and M. polynoda

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Appendix 2. Perennial grass species with potential for domestication A. Native species Species Attributes Source from

Appendix 1A

Astrebla spp. (Mitchell grasses – A. lappacea, A. pectinata)

Seed gathered, ground and made into damper by Aboriginal people, relatively shatter resistant.

19, 25, 37, 47

Brachiara spp. (millet) Used by the Pitjantjatjara and Jankuntjatjara people 37, 77

Brachiara miliiformis

(armgrass millet)

One of the larger grass seeds used by Aboriginal people – 2 mm diameter, 3.2 mg in weight, 1000–2000 seeds per plant (2.3 g seed/plant)

14, 37, 63

Dactyloctenium radulans (button grass)

Found adjacent to water in the Tanami and Great Sandy deserts, seeds used for human consumption

14, 63, 77

Danthonia spp. (wallaby grass, white-top)

High forage value 24, 47, 73

Eragrostis eriopoda (woollybutt, wire wanderrie grass, neverfail, wangunu, atjira, naked woollybutt)

Perennial, small-seeded spikelets, easy to husk, easy to grind, used to make bread or damper, seed ripening in autumn or winter after summer rains. Seeds 0.12 mg, 10 000–20 000 seeds per plant (1–23 g seed/plant)

12, 14, 19, 25, 36, 37, 49, 63

Eragrostis laniflora 14

Microlaena stipoides (weeping grass, meadow rice grass, Ehrharta stipoides)

Selected for seed yield by research groups interested in its forage potential

47, 95

Panicum australiense (bunch panic)

Used by Aboriginal people; smaller plant than P. decompositum (an annual), seeds usually gathered from the ground

14, 49, 63

Panicum decompositum (Australian millet, umbrella grass, native millet, tindil, coolly, windmill grass, papa grass, kaltu kaltu, altjurta)

Aborigines in the Murray Darling ground the small grains into flour and roasted or baked into damper, seed ripening after summer rains (annual or perennial)

3, 14, 19, 25, 29, 36, 37, 49, 54, 63

Sporobolus (yakka grass, fairy grass (S. actinocladus — Qld, NT, S. caroli — all states))

Very small grains, Aborigines ground the grains between stones and baked a type of damper in the ashes

25

Tetrarhenia leavis (native WA rice grass)

Large seeded native grass, medium to high rainfall (John S Pate, pers. comm.)

Themeda triandra (kangaroo grass, red grass, red-oat, red-oat grass, Themeda australis?)

Perennial, grain eaten during famine, used for fodder/paper, Africa/South Africa/Tropical Asia/Temperate Asia/Australasia

25, 36, 47

Whiteochloa cymbiformis (Panicum cymbiforme)

Australasia 14

Yakirra australiensis (bunch panic grass)

Australia (Michael O’Connor, pers. comm.)

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B. Exotic species Species Attributes Source from

Appendix 1B

Agropyron elongatum (perennial relative of wheat)

26

Agropyron ponticum (perennial relative of wheat)

26

A. ponticum x T. aestivum (perennial wheat)

Crosses made by Sherman and Sandos at University of California, Davis, 1950s

(Stephen Jones, pers. comm.)

Bromus mango (Bromus burkartii)

Annual (?) Argentina, Chile, reputed to have been cultivated for grain in prehistoric times

(Mark Blumler, pers. comm.)

Bromus stamineus (grazing brome)

Perennial, large seeds, origin Chile, Argentina (Mark Blumler, pers. comm.)

Bromus wildenowii (prairie grass, rescue grass)

Short-lived perennial, large seeds, well-drained soils, pH>6, South America/Argentina

(Mark Blumler, pers. comm.)

Dactyloctenium spp. 10

Echinochloa colonum (shama millet, corn panic grass, deccan grass, jungle ricegrass)

Eaten as grain in dry years, likes the wet, tropics/subtropics/warm temperate regions

5

Echinochloa pyramidalis? (antelope grass, limpopo grass)

Fodder, flour, Africa/South Africa 5

Echinochloa stagnina (bourgou, burgu grass, hippo grass, long-awn water grass)

Floodplains, mainly for fodder, seed has been harvested for food, produces sugar for beverages, Africa/South Africa/Tropical Asia

5

Elytrigia intermedia (intermediate wheat grass, wild triga, Agropyron intermedium, Thinopyrum intermedium, Elymus hispidus, Agropyron glaucum)

Originally from Temperate Asia/Tropical Asia/Europe, potential to become a commercially viable crop, 400 lines evaluated.

24, 25, 26, 28

Elytrigia pontica (tall wheat grass, Thinopyrum ponticum)

~350 mm rainfall, Temperate Asia/Europe (Doug Johnson, pers. comm.)

Eragrostis spp. 6, 10

Eragrostis gangetica? Grains used as a famine food 5

Leymus racemosus (mammoth wild rye)

Temperate Asia/Europe 19

Panicum staginum Grain no good, produces thick syrup useful for confections and sweet beverages

5

Panicum subalbidum? 5

Panicum turgidum (afezu, merkba, desert grass, markuba, taman, tuman)

Deep-rooted, drought-resistant (30–250 mm), large seeds, palatable (not suitable for bread), grains used mainly for porridge, Africa/Temperate Asia/Tropical Asia

5, 10

Setaria sphacelate (Panicum sphacelatum)

Robust, seeds eaten as a famine food, used for hay/silage/grazing, Africa/Temperate Asia

5

Sporobolus fimbriatus? Africa 5

Stenotaphrum dimidiatum (Panicum sphacelatum)

Stoloniferous perennial, Africa/Tropical Asia 5

Stipagrostis pungens (drinn, Aristida pungens)

Relatively tall (to 1.5 m), black grains, deep roots, drought resistant, Africa/Temperate Asia

5, 10

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Species Attributes Source from Appendix 1B

Tripsacum dactyloides (eastern gama grass)

North & South America 12

Urochloa brizantha (Brachiaria brizantha, bread grass, palisade grass, signal grass)

Africa/South Africa 5

Urochloa mosambicensis? (sabi grass)

Grains boiled, Africa/South Africa/Tropical Asia 5

Urochloa trichopus? (Panicum trichopsis)

Grains sometimes eaten

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Appendix 3. Networks Tim Barden Ko-warra Transplants PO Box 396 Moama NSW 2731 kowarra@mcmedia.com.au Avril Baxter Conservation and Land Management PO Box 100 Narrogin WA 6312 avrilb@calm.wa.gov.au Wallace and Sue Binnie Farmers RMB 1130 Alexanders Road Bungeet Vic 3726 Wendy Bradshaw Bushcare Support Ofiicer Murray Wells Road Tambellup WA 6320 wendyb@gawa.comdek.net.au Roy Butler Agriculture WA PO Box 432 Merredin WA 6415 rbutler@agric.wa.gov.au Ian Chivers Managing Director Native Seeds P/L PO Box 133 Sandringham Vic 3191 racsols@nativeseeds.com.au Ian Cole Ecologist Department of Infrastructure, Planning and Natural Resources Centre for Natural Resources PO Box 445 Cowra NSW 2794 ian.cole@dipnr.nsw.gov.au Dr Stan Cox Senior Research Scientist The Land Institute 2440 E. Water Well Road Salina Kansas 67401 USA cox@landinstitute.org

Dr Kingsley Dixon Director, Science Kings Park and Botanic Gardens West Perth WA 6005 kdixon@kpbg.wa.gov.au Judi Earl Australian Information and Monitoring Services Armidale NSW 2350 judiearl@bigpond.com Chris Findlay Head, Revegetation Operations Native Seeds P/L PO Box 133 Sandringham Vic 3191 chrisf@nativeseeds.com.au FloraBase www.florabase.calm.wa.gov.au Lindsay and Margaret Gentle Farmers Tamworth NSW 2340 lvgentle@northnet.com.au Sascha Groeneweg SARDI Genetic Resource Centre Waite Campus GPO Box 397 Adelaide SA 5001 groeneweg.sascha2@saugov.sa.gov.au David Harris Chemistry Centre 125 Hay Street East Perth WA 6004 dharris@ccwa.wa.gov.au Professor Robert Henry Director CRC for Innovative Grain Food Products c/– Centre for Plant Conservation Genetics Southern Cross University Military Road PO Box 157 Lismore NSW 2480 rhenry@scu.edu.au

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Dr Wes Jackson President The Land Institute 2440 E. Water Well Road Salina Kansas 67401 USA jackson@landinstitute.org Doug Johnson USDA-ARS Forage and Range Research Laboratory Utah State University Logan UT 84322-6300 USA daj@cc.usu.edu Christine Jones Department of Land and Water Conservation PO Box 199a Armidale NSW 2350 cjones@dlwc.nsw.gov.au Assoc. Prof. Stephen Jones Crop and Soil Sciences Washington State University PO Box 646420 Pullman WA 99164-6420 USA joness@wsu.edu Des Lang Department of Infrastructure, Planning and Natural Resources Research Centre PO Box 462 Gunnedah NSW 2380 des.lang@dipnr.nsw.gov.au Peter Latz PO Box 2482 Alice Springs NT 0871 Alec Lazenby CSIRO Plant Industry GPO Box 1600 Canberra ACT 2601 alec.lazenby@csiro.au Assoc. Prof. Slade Lee Centre for Plant Conservation Genetics Southern Cross University Military Road PO Box 157 Lismore NSW 2480 slee@scu.edu.au Greg Lodge Tamworth Centre for Crop Improvement NSW Agriculture RMB 444 Tamworth NSW 2340 greg.lodge@agric.nsw.gov.au

Dallas Lynch Bushcare Support Centre/ Western Australian Native Grasses Society (WANGS) c/– Avon Catchment Network PO Box 311 Northam WA 6401 dlynch@agric.wa.gov.au Dr Neville Marchant Director, WA Herbarium Conservation and Land Management Locked Bag 29 Bentley Delivery Centre Bentley WA 6983 nevillem@calm.wa.gov.au Terry MacFarlane Conservation and Land Management Brain Street Manjimup WA 6258 terrym@calm.wa.gov.au Meredith Mitchell Research Officer Rutherglen Research Centre Department of Primary Industries RMB 1145 Chiltern Valley Road Rutherglen Vic 3685 Meredith.Mitchell@dpi.vic.gov.au Bill Mollison Permaculture Institute 31 Rulla Road Sisters Creek Tas 7325 tagariadmin@southcom.com.au Geoff Moore Department of Agriculture Locked Bag 4 Bentley Delivery Centre Bentley WA 6983 gmoore@agric.wa.gov.au Michelle Murphy Department of Botany University of New England Armidale NSW 2351 mmurphy2@metz.une.edu.au Robert Myers Native Grasses Resource Group PO Box 250 Birdwood SA 5234 www.mlrcp.sa.gov.au/native_grass/index.html

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Zhongnan Nie Pastoral and Veterinary Institute Mount Napier Road Private Bag 105 Hamilton Vic 3300 zhongnan.nie@dpi.vic.gov.au Michael O’Connor CSIRO Sustainable Ecosystems Private Bag 5 Wembley WA 6913 Michael.O’Connor@csiro.au Rex Oram CSIRO Plant Industry PO Box 1600 Canberra ACT 2601 rex.oram@csiro.au Tony and Lindy Piggins Farmers ‘Bilawi’ Corowa NSW 2646 lindy@dragnet.com.au Mike Quarmby Reedy Creek Nursery Princes Highway Reedy Creek SA 5275 Alistair and Jane Robb Farmers ‘Burraja’ Corowa NSW 2646 Paul Sanford Department of Agriculture 444 Albany Highway Albany WA 6330 psanford@agric.wa.gov.au Brian Sindel CSIRO Plant Industry PO Box 1600 Canberra ACT 2601 Brian.Sindel@csiro.au Dr Jesse Skoss Palo Verde Plant Research Resource Association RMB 28 Manjimup WA 6258

Prof. German Spangenberg Research Director Plant Genetics and Genomics Primary Industries Research Victoria Department of Primary Industries Victoria german.spangenberg@dpi.vic.gov.au STIPA (based in NSW) http://home.winsoft.net.au/stipa/ Peter Stone Farming Systems Specialist CSIRO Sustainable Ecosystems Private Bag 5 Wembley WA 6913 Peter.Stone@csiro.au Zoe Toll Agronomist Native Seeds P/L PO Box 133 Sandringham Vic 3191 tolly3566@fastmail.fm Darren Vincent Turf and Lawn Care Wangaratta Vic 3676 darren@turfandlawncare.com.au Cathy Waters NSW Agriculture Australian Native Grasses and Legume Seed Industry Association PMB 19 Trangie NSW 2823 Cathy.waters@agric.nsw.gov.au Dr Colin Wellings Senior Research Scientist NSW Agriculture Plant Breeding Institute Private Bag 11 Camden NSW 2570 colinw@camden.usyd.edu.au Professor Wal Whalley Honorary Fellow, Botany School of Environmental Sciences and Natural Resources Management Faculty of the Sciences University of New England Armidale NSW 2351 rwhalley@metz.une.edu.au