Leguminosae

Common name
Lotononis

Origin and geographical distribution

Occurs naturally in South Africa (Transvaal), Mozambique, Namibia, Swaziland and Botswana between latitudes 20°S to 26°S (Strickland, 1972; Van Wyk, 1991). It has been introduced to the subtropical, tropical, temperate and Mediterranean regions of all continents. The earliest reference to evaluation outside its native habitat is in nurseries at Kitale, Kenya in 1951 (Bogdan, 1955). This species was collected in 1952 by J.F. Miles (Australian Plant Introduction Officer) at Worcester Veldt Reserve (Bryan, 1961), also outside its native habitat, therefore some evaluations of L. bainesii pre-dated the 1950s.

Description

Herbaceous perennial with slender stoloniferous stems. Leaves palmate, mostly trifoliate and rarely with 4 or 5 leaflets. There is usually 1 leaf per node. Leaflets are polymorphic: linear, ovate, obovate or lanceolate. The central leaflet being up to 3-cm wide and 6-cm long while the other 2 leaflets are smaller. Petioles are 6-mm to 5-cm long. Fully developed leaves are generally glabrous, giving a shinny appearance to the foliage. A taproot is developed as well as numerous secondary roots and similar root systems develop from the nodes of the stolons. Inflorescences are racemes, which may be contracted to dense umbellate heads with few to more than 20 florets, the peduncles up to 25-cm long. Florets small, yellow; calyx funnel-shaped (4 mm to 5 mm), slightly silky, with very short subdeltoid teeth, corolla up to 14 mm, standard and keel almost equal, the wings small and shorter. Pods linear-oblong, 8 to 12 mm long, many-seeded, (up to 20), and tardily dehiscent. Seeds cream-yellow to magenta-rose, obovoid and asymmetrically heart-shaped, the cotyledonary lobe larger than the radicle, laterally compressed (Oram, 1990; Van Wyk, 1991; Real et al., 2004; Martínez and Risso, 2004).

Season of growth

Its native environment is dry and cold in winter and it is not adapted to grow at that season and is a spring/summer-growing perennial. If sown in a winter growing environment, it will grow slowly in winter, mainly limited by day-light and temperature. In most climates the best growing seasons are spring and summer; however, summer production is highly correlated with moisture availability.

Frost tolerance and regrowth

Lotononis bainesii is quite frost-tolerant compared to many subtropical/tropical legumes (Gates, 1973); its winter growth is minimal, but it remains green after minor frosts of -1 °C to -3 °C, while after frosts of -4 °C to -10 °C, green foliage is burnt and turns black. However, after a few days of warm weather in winter, plants regrow both from stolons and the crown. In subsequent frosts, new growth may be burnt again, but plants survive. In spring, after the frost, established plants recover well.

Lotononis bainesii appears to be a long-day plant which flowers mainly in the spring, with a secondary flowering in late summer (Bryan, 1961; Wright, 1964). However, in a second year or older crop, L. bainesii starts flowering as soon as it begins growth after winter and keeps flowering through spring, summer and autumn. This indeterminate flowering characteristic makes it difficult to decide the optimum harvesting time, and site-specific research is necessary (Real et al., 2005). It is very sensitive to light and will not tolerate excessive shading (Bryan et al., 1971).

Lotononis bainesii is drought tolerant, more so than Desmodium intortum and almost as much as Macroptilium atropurpureum. In severe droughts leaves may dry, but plants are able to regrow from reserves in the tap-root. After very severe drought it has been known to regenerate from seeds.

Lotononis bainesii is small seeded and the early growth is slow. If there is a minor drought during establishment, strategic irrigation may be beneficial to allow plants to establish. After the establishment phase, plants are very drought tolerant and irrigation should not be needed except in seed production schemes.

Lotononis bainesii is tolerant of temporary water-logging. Wright (1964) reported it survived natural floods of up to six days. Whiteman et al. (1984) and Javier (1985) reported a water-logging pot-experiment in which pots were continuously flooded to 5 cm above the soil surface for 10 and 21 days. Adaptation to flooding (i.e. tolerance) was related to rapid production of adventitious roots from immersed stems and branches. Yield of waterlogged plants were 90% of the drained control.

Bryan (1961) reported that L. bainesii requires at least 875 mm annual rainfall and has been most successful between 1,125 and 1,625 mm. However, recent introductions of this species into Mediterranean regions with 550 mm of annual rainfall concentrated during winter have been able to persist for more than 10 years (pers. comm. Howieson and Yates, 2005). In its native environment it is found in areas with rainfall as low as 350 mm (Strickland, 1972).

Lotononis bainesii grows well on a wide range of soil types from shallow stony ones to deep sands. Bryan (1961) suggested that is best adapted to sandy soils such as lateritic podzolic, low-humic gley, red-yellow podzolic and meadow podzolic soils. However, it will grow on heavy clay (Wright, 1964). In sand culture, it has grown and nodulated quite successfully at pH 4.0 (Andrew and Johnson, 1976). It also tolerates an excess of aluminium and manganese better than most subtropical legumes (Cameron, 1985); however, high levels of aluminium are detrimental for the species.

Norris (1958) reported that L. bainesii requires an extremely specific type of root-nodule bacteria which develops red colonies on culture medium. At that time, Norris thought that the bacteria were Rhizobium. However, Jaftha et al. (2002) concluded, partially on the basis of 16SrRNA sequences, that the bacteria were not related to any rhizobial group and in fact, they were closely related to Methylobacterium nodulans. On the basis of current studies at the Centre for Rhizobium Studies, Perth, Western Australia, Methylobacterium strains which nodulate and fix nitrogen with L. bainesii do so with the closely related species L. listii, but not with L. angolensis. All three Lotononis species are in the Listia subsection. Up to 2005, the exact species of bacterium within the Methylobacterium genus that nodulates L. bainesii has not been identified, although it is considered unlikely to be M. nodulans but a closely related species

Where pasture is kept short, L. bainesii can spread naturally. Wright (1964) reported that it invaded the tall Paspalum urvillei pasture at Coolum in south-eastern Queensland. Seed is also spread in the dung of cattle that graze the plant.

Lotononis bainesii will only establish satisfactorily if sown on a fine, well-prepared compact seed bed. Seed is so small that it needs a weed-free planting medium. Usually an application of herbicide to control all existing vegetation is beneficial for optimal establishment.

Seed is usually broadcast on the surface, or with a minimum soil disturbance. If a roller is available, the seed bed can be rolled both before and after planting; otherwise, a very light harrowing is necessary. The plant will also establish readily from cuttings and turves. A turf 15 cm square is suitable planting material, each piece being placed about 2.5 m apart in rows 2 m apart, to allow inter-row cultivation (Wright, 1964). Bryan (1968) found that oversowing failed in Paspalum commersonii pasture. From 2003 to 2005, several oversowings conducted in Uruguay also failed in pastures dominated by Paspalum notatum and Axonopus affinis; there also, recent research shows the need to reduce competition effectively from the existing canopy, i. e. application of a total herbicide in a dose that may temporarily stop growth (Risso et al, Unpublished).

Blumenthal and Hilder (1989) conducted several pot experiments with seeds sown on the surface and at 5, 10, 25 and 50 mm of depth. Seedlings were not able to emerge if planted deeper than 10 mm. All irrigation treatments that allowed continuous moisture in the first centimetre of soil were optimum for germination.

Seeds are small and seedling vigour is poor, therefore when establishing L. bainesii, it is necessary to provide the best sowing conditions. For optimum growth at the seedling stage, high quality seed needs to be sown at optimum soil temperature and humidity in a weed free environment. McDonald (2002) studied germination from 8 to 44 ºC and found that between 16 to 36 ºC germination was at least 67 % with a maximum germination of 99.2 % at 28 ºC. With temperatures below 16 ºC or above 36ºC, germination dropped to less than 15 %. For dry environments, seed needs to be sown at least 3 months before a dry season starts, so that the plants are well established and with a minimum root-depth. These parameters need to be considered when determining the best sowing time for each region. Sowing rate between 0.5 kg/ha (140-160 seeds/m²) to 3.5 kg/ha (980-1,155 seeds/m²) have been used. However, with as few as 10 to 20 established plants per m², full coverage of the area can be expected. Recommended seed rate should be based on achieving no fewer than 10 well established plants/m² and take into consideration time of sowing (expected temperature and rainfall), seed bed preparation and seed price to decide the seed rate.

Thousand seed weight (TSW) of viable seed can vary from 0.2 g (5,000,000 seeds/kg) to 0.45 g (2,200,000 seeds/kg). Seed of TSW of less that 0.2 g is mainly empty seed with very poor germination and vigour (Real et al., 2005). A commercial seed lot would normally have a TSW of 0.3 g to 0.35 g.

The percentage of hard seed is fairly high, in heavy and high quality seed, of up to 80 %. Light and low-quality seed usually have a lower percentage of hard-seed (Altuve et al. 1991; Real et al., 2005). Purple and brown seed keep their viability better than yellow and green seed (Poulsen, 1966).

To break dormancy, scarify L. bainesii with fine sand paper, or scarifying machines for large seed quantities. Scarification intensity should be adjusted to avoid seed damage. It can also be treated with concentrated sulphuric acid (sp. gr. 1.8) for 20 minutes, wash and dry (Prodonoff, 1968) or hot water between 20 and 30 °C to improve germination (Poulsen, 1966; Altuve et al., 1991). However, water temperatures above 35 °C have been found to lower germination. Inoculation with the appropriate Methylobacterium strain is absolutely necessary.

Lotononis bainesii responds to fertilizer, although it has moderate demands for phosphorus. On light sandy soil, it should be given a complete mixture of fertilizer.

Norris (1956) has shown that L. bainesii nodulated successfully in sand culture at pH 4.0 and in the presence of only 5 ppm of calcium. Lotononis bainesii calcium uptake is 1.1 to 1.6 % of the dry matter (Andrew and Hegarty, 1969).

Andrew and Robins (1969a) determined the critical percentage of phosphorus in the top growth of L. bainesii at the immediately pre-flowering period as 0.17 %. It was shown to be almost as effective as Stylosanthes humilis in extracting its phosphorus needs from low-fertility soils. Andrew and Robins (1969b) further showed that L. bainesii and S. humilis were the most effective in producing nitrogen per unit mass of soil, especially on low phosphorus soils. The percentage of nitrogen in the tops of L. bainesii increased from 3.29 % with 125 kg/ha superphosphate to 3.46 % at 250 kg/ha and 3.91 % at 1000 kg/ha. Risso et al. (2004) conducted a preliminary fertilization experiment in Uruguay with increasing levels of P₂O₅ (0, 30, 45, 60, and 120 kg/ha). There was a large increase in dry matter yield for the first 30-45 kg of P₂O₅/ha in comparison with the un-fertilized control. Above that amount, in spite of some increase in its proportion in the total canopy and to some extent in volume of dry matter, increases were minor, confirming the phosphorous efficiency of this legume.

Andrew and Robins (1969c) determined the critical percentage for potash in the plant tops of L. bainesii as 0.90 %. They found that plants yielded only 23 % of their maximum yield without the addition of potash, and required more potash than Siratro. Lotononis bainesii and Desmodium intortum are usually high in potassium (Andrew and Robins, 1969d). Lotononis bainesii growing in a soil with adequate potash had a potassium concentration of 1.08 g DM/kg, while a deficient plant recorded only 0.41 g DM/kg in the tops. The initial effect of potassium deficiency in L. bainesii is the appearance of a generalized interveinal chlorosis of the leaflets of the older leaves, commencing as a blotching of the leaflets, particularly toward the tips and margins. With increasing deficiency, the chlorosis intensifies, particularly toward the tips of the leaflets, which finally develop light brown, irregular shaped necrotic spots which enlarge and coalesce to form large brown areas with some distortion around the necrotic areas. Finally, necrosis encompasses the entire leaflet and death ensues. The petioles remain unaffected for some time but eventually abscise, and even the stipules of the affected petioles develop necrosis. In general, potassium deficiency symptoms are more severe on the central upright stem of the plant than on the lateral stems. Under conditions of extreme deficiency, even the freshly expanding leaves are chlorotic and malformed and have pinched tips (Andrew and Pieters, 1970a).

Andrew and Robins (1969b) found that the sodium content of L. bainesii was high in comparison to other tropical legumes and increased with fertilization with sodium dihydrogen phosphate at the expense of potassium.

Manganese - the initial effects of manganese toxicity are interveinal chlorosis in the young leaves and chlorotic shoots. With increasing toxicity, irregularly shaped brown areas occur on both leaflet surfaces; these subsequently become necrotic. At the same time, younger expanding leaves are small, chlorotic and malformed exhibiting unusual leaflet shape, distortion of the edges, and intense puckering of the central portion of the leaflet. This is accompanied by outward curvature of the leaflet margins and a downward curving of the distal portion of the leaflets. Brown-coloured necrotic areas occur on the petiole and stems. Restricted growth gives rise to strong secondary growth which is chlorotic at emergence (Andrew and Pieters, 1970b). Lotononis bainesii is tolerant of excess manganese and the toxicity threshold value is 1320 ppm (Andrew and Hegarty, 1969).

Lotononis bainesii combines well with Paspalum commersonii, P. plicatulum and P. dilatatum, and especially well with Digitaria decumbens (pangolagrass), as these species can withstand heavy grazing. It will grow with tall grasses such as Setaria anceps if they are defoliated frequently. It is less compatible with thickly matted Kikuyu grass (Pennisetum clandestinum). It is compatible with Macroptilium lathyroides, Trifolium repens and Centrosema pubescens (Bryan, 1961). The practise of sowing a winter-active crop such as annual ryegrass, oats, triticale, etc. into an established L. bainesii sward of two or more years old helps to protect the legume from severe frosts, keeps the area relatively weed free and during the least productive season for L. bainesii the land can be used productively. The grass crop should be established without nitrogen-based fertilisers as nitrogen should be readily available from that provided by L. bainesii in the previous season. The grass should have a short growing season so that it can be removed by cutting, grazing, harvesting or by spraying a grass herbicide to the area when L. bainesii starts to grow after winter.

Lotononis bainesii in the seedling stage is short and thus susceptible to shading by tall weeds. Weeds might also be a problem for L. bainesii seed crops if their seeds are of a similar size.

The only two broadleaf herbicides that can be recommended for L. bainesii (from information generated up to 2004) are Flumetsulan and Imazethapyr at doses recommended for pasture legumes in the country where they are to be sprayed. Grass herbicides can be used as normally recommended for forage legumes (Rios, 2004).

Seeds of L. bainesii are small and even though the relative growth rate of seedlings is high (Blumenthal and Hilder, 1989), their growth is not vigorous enough to compete with weeds and to establish a deep root system to avoid losses due to surface drought. Therefore, seed quality, time of sowing and sowing conditions should be optimized.

Lotononis bainesii fixed an average of 200 kg. N/ha over 5 years at Samford, south-eastern Queensland. Bryan (1961) showed that its addition to a pasture of Paspalum commersonii and P. plicatulum increased the nitrogen content of the grass by 14 %. On the basis of studies at the Centre for Rhizobium Studies, Perth, Western Australia, the Methylobacterium strains which nodulate and fix nitrogen with L. bainesii are highly persistent in acid and sandy soils and can nodulate well to fix nitrogen in stressful environments. The Methylobacterium strains appear to colonize poor soils well and can spread to nodulate stoloniferous roots developed on L. bainesii.

New stolon development accompanies grazing and so L. bainesii is able to withstand some grazing pressure. Jones et al. (1967) and Whiteman (1969) found that it did not persist with high yields for more than a few years, but Bryan (1968) found it more persistent at Beetwah in south-eastern Queensland. If a L. bainesii stand is defoliated continuously or at frequent intervals, it adopts a prostrate habit that is very beneficial for the persistence of the species. An experimental sward of L. bainesii survived sheep grazing for 10 years at Jerdacuttup, Western Australia.

Lotononis bainesii is stoloniferous and roots from the nodes. However, stolons will only be close to the ground if there is no plant competition. If there is competition, stolons will grow over the companion plants impeding contact of the new roots with the soil so they will not, therefore, develop further. In a competitive environment, L. bainesii plants will only have the original seedling tap-root. To encourage runners to root down, L. bainesii should be kept fairly closely grazed to remove competition by other species. If the plant is only living from its original seedling tap-root, that root gets old and diseases can attack and kill the plant. If, however, the plant is continuously rooting at the nodes, it is always regenerating itself, always young, and very tolerant to diseases and mismanagement. This is why heavy grazing (within reason) improves its persistence and is beneficial to the success of the species. Risso et al. (2004) studied the response to defoliation when the sward (L. bainesii and Uruguayan native grassland) reached 10 or 15 cm of height and cut to a height of 5 or 3 cm. The best forage yield (6,700 kg/ha) was obtained when the sward was allowed to grow to 15 cm of height and cut to a 5 cm stubble height.

Lotononis bainesii recovers readily after fire.

Lotononis bainesii is a tetraploid with 2n = 4 x = 36 (Byth, 1964). Flowers were reported to be cleistogamous and therefore, self-pollinated (Hutton, 1960, Byth, 1964). However, recent evidence suggests that the species is chasmogamous and in fact highly cross-pollinated, needing pollinators to set seed (Real et al. 2004, Dalla Rizza et al. 2004, Real and Altier, 2005, Real et al., 2005).

At Rodd's Bay, Queensland, 1,850 kg DM/ha of L. bainesii were harvested after four months' growth. At Beerwah, the yield was 5,050 kg DM/ha year. In grazing trials at Beerwah with 6 weeks rest between grazings, yields have ranged from 25 kg/ha in a dry early spring to 2673 kg DM/ha in mid-summer. With pangolagrass (Digitaria decumbens) and 8 weeks between grazings and a stocking rate of 1 beast/0.80 ha, L. bainesii yield range was between 250 to 1,912 kg/ha while the pangolagrass yield ranged between 1,564 to 2,340 kg/ha (Bryan, 1961). At Samford, Jones et al. (1967) harvested over a four year period 3,850 kg/ha, 2,937 kg/ha, 3,751 kg/ha and 1,078 kg/ha dry matter yield, respectively, of the L. bainesii component within a L. bainesii-grass association. This dry matter yield represented 33.5, 43.3, 27.9 and 12.9 % respectively, of the total dry matter yield of the sward. Under irrigation at Perth, Western Australia, in a Mediterranean climate (lat. 32°S), Roberts and Carbon (1969) obtained average annual yields at two sites of 8,268 kg/ha. In a subtropical climate in Uruguay, establishment year yield was 5,000 kg DM/ha and 7,000 kg DM/ha, respectively, 5 months after planting in 2003 and in 2004.

No hay appears to have been made with L. bainesii, but if care is taken to preserve the leaf, hay should be of high quality. Catchpoole (1970) made excellent silage from L. bainesii cut at 2.5 to 4 cm above ground level without the addition of molasses. The pH of the silage was below 4.2 with a low content of volatile constituents, and small losses of dry matter and nitrogen during storage. The material was chaffed for ensilage. Fermentation followed the usual lactic acid sequence, but development of lactic acid and attainment of the final pH were slow. The final silage had 25.6 % dry matter with 15.37 % protein on a dry-matter basis.

Lotononis bainesii is an excellent species for deferred use because it does not turn woody and is continuously renewing its foliage through its growing points. However, large accumulations of forage might be detrimental to its persistence as explained in the defoliation section. If the growing region has a shortage of forage during autumn (autumn gap) and rains in summer, this species is especially good. If the shortage is in winter, because of its greenness and tolerance to mild frosts, it can be very valuable as deferred forage. If frosts are severe, the foliage is burnt and turned to a black colour, disappearing into the ground after a few rain events, therefore not suitable to be deferred for winter grazing in these cold regions.

Feeding value

Chemical analysis:
The results of 14 analyses showed that the crude protein content of the dry matter reached 25.9% (Bryan, 1961), with a mean value of 18.3 %. The full analyses obtained by Milford (unpublished) were: crude protein 19.3 %, crude fibre 27 %, ether extract 4%, nitrogen-free extract 41.6%, ash 8.1% and moisture 74%. Milford (1967), reported the feeding value of L. bainesii, as being the highest in comparison with Macroptilium lathyroides, Macroptilium atropurpureum, Desmodium uncinatum, Vigna vexillata, Stylosanthes guianensis and Crotalaria lanceolata. Chemical analysis of 30 samples during 2002, 2003 and 2004, had the following mean values: digestibility of organic matter 61%, crude protein 15%, acid detergent fibre 31%, neutral detergent fibre 52% and ash 7%.

Cattle and sheep (after a short period of getting familiar with the new fodder) graze L. bainesii eagerly. It was reported by Wright (1964) to be highly palatable.

The tannin content measured with the vanillin test is 0.6% for inflorescences, 0.7% for leaves and 0.4% for stems. These might be why L. bainesii is considered a non-bloating legume. Levels are low enough not to interfere with palatability and might protect protein degradation in the rumen.

No toxicity has been observed with L. bainesii (Wright, 1964; Cameron, 1985), although the lush growth will cause a temporary taint in milk of dairy cows grazing it. Both beef cattle and sheep have been grazing L. bainesii in different countries and situations, and no ill effects have been reported up to 2005.

The seed crop is ready for harvesting when about 80 % of the crop has purple seed (Altuve et al, 1991). This will usually contain 50 % fully mature purple seed, 30 % soft purple seed and 20 % green, yellow and brown seed. The crop should be mowed to 5 cm of height with a disc mower and left in the swath for 48 to 72 hours to dry. Once the material has reached 80 % DM it is then gathered with a combine harvester with the cylinder set at 1200 rpm, 2-mm cylinder-concave separation and a concave for forage seed with 2.5 mm of separation between bars. To avoid any seed loss in the field, the air draft should be reduced to a minimum (Martínez and Risso, 2004; Real et al., 2005).

Seed production of L. bainesii at a commercial scale was about 30 to 55 kg/ha. (Bryan 1961) and could be as high as 262 kg/ha (Altuve et al. 1991). Real et al. (2005) studied the weekly seed production of the species. In 13 weekly harvests, it accumulated a total of 656 kg/ha, indicating the potential of seed production for the species. Due to pod shattering, successive flushes of pods could not be accumulated for harvesting. The maximum weekly production was of 267 kg/ha, indicating the potential harvestable seed. Cameron (1985) reported seed yields of about 300 to 400 kg/ha standing in the paddock, but harvested seed of only 20 kg/ha.

A minimum of 50 % germination and 93 % purity with a maximum of 45 % hard seed is required in Australia. In Uruguay, it is also 93% purity, 7% seeds of other crops, 1% weeds, 7% inert matter and a minimum of 50 % germination. The seed is germinated at 20 to 30°C.

‘Miles’, released by CSIRO, Australia, 1966

‘Beit’, Zimbabwe, 1972

‘INIA Glencoe’, released by INIA, Uruguay, 2003.

Cook et al. (2005) reported several fungal diseases such as leaf spot (Cercospora sp.), Sclerotium rolfsii, Fusarium and Pythium root and stolon rots, flower blight (Botrytis sp.) and Rhizoctonia solani. L. bainesii is very susceptible to legume little-leaf disease caused by a phytoplasma carried by the common brown leafhopper (Orosius argentatus: Cicadellidae), more so if it is grazed lightly in late summer. Bean yellow mosaic virus (BYMV, Potyvirus) spread by over 20 species of aphids can also be a problem. Grazing helps in minimizing most disease damage. It also is attacked by the spiral (Helicotylenchus dihystera) and root-knot (Meloidogyne hapla) nematodes. Main insect pests are heliothis (Helicoverpa armigera) and pod-sucking bugs such as Riptortus serripes (Alydidae) and Nezara viridula (Pentatomidae), which can be a problem in seed stands. It is resistant to amnemus weevil (Amnemus quadrituberculatus).

Main references

Altuve, S.M., Ramirez, M.A., Perez, D. and Royo Pallares, O. (1991) Lotononis bainesii, una leguminosa para el subtrópico húmedo de Argentina. 2. Producción de semillas. In: XII Reunião do grupo técnico regional do cone sul (Zona campos) em melhoramento e utilização de recursos forrageiros das areas tropical e subtropical. Bagé, RS, Brasil. p. 91-94.

Andrew, C.S. and Hegarty, M.P. (1969). Comparative responses to manganese excess of eight tropical and four temperate pasture legume species. Australian Journal of Agricultural Research, 20: 687-696.

Andrew, C.S. and Johnson, A.D. (1976). Effect of Calcium, pH and Nitrogen on the growth and chemical composition of some Tropical and Temperate Pasture Legumes. II. Chemical composition (Calcium, Nitrogen, Potassium, Magnesium, Sodium and Phosphorus). Australian Journal of Agricultural Research, 27: 625-636.

Andrew, C.S. and Pieters, W.H.J. (1970a). Effect of potassium on the growth and chemical composition of some pasture legumes. III. Deficiency symptoms of 10 tropical pasture legumes. Melbourne, CSIRO Australian Division of Tropical Pastures. Tech. Paper No. 5.

Andrew, C.S. and Pieters, W.H.J. (1970b). Manganese toxicity symptoms of one temperate and seven tropical pasture legumes. Melbourne, CSIRO Australian Division of Tropical Pastures. Tech. Paper No. 4.

Andrew, C.S. and Robins, M.F. (1969a). The effect of phosphorus on the growth and chemical composition of some tropical pasture legumes. I. Growth and critical percentage of phosphorus. Australian Journal of Agricultural Research, 20: 665-674.

Andrew, C.S. and Robins, M.F. (1969b). The effect of phosphorus on the growth and chemical composition of some tropical pasture legumes. II. Nitrogen, calcium, Magnesium, potassium and sodium contents. Australian Journal of Agricultural Research, 20: 675-685.

Andrew, C.S. and Robins, M.F. (1969c). The effect of potassium on the growth and chemical composition of some tropical pasture legumes. I. Growth and critical potassium percentages. Australian Journal of Agricultural Research, 20: 999-1007.

Andrew, C.S. and Robins, M.F. (1969d). The effect of potassium on the growth and chemical composition of some tropical pasture legumes. II. Potassium, calcium, Magnesium, sodium, nitrogen, phosphorus and chloride. Australian Journal of Agricultural Research, 20: 1009-1021.

Blumenthal, M.J. and Hilder, T.B. (1989). Emergence and early growth of Lotononis bainesii cv. Miles on a cracking soil compared with four other tropical legumes. Australian Journal of Experimental Agriculture 29: 193-199.

Bogdan, A.V. (1955). Herbage plants at the Grassland Research Station, Kitale, Kenya. The East African Agricultural Journal, 151-165.

Bryan, W.W. (1961). Lotononis bainesii Baker a legume for subtropical pastures. Australian Journal of Experimental Agriculture and Animal Husbandry. Vol 1: 4-10.

Bryan, W.W. (1968). Grazing trials on the Wallum of southeastern Queensland. II. Complex mixtures under common grazing. Australian Journal of Experimental Agriculture and Animal Husbandry. Vol 8: 683-690.

Bryan, W.W., Sharpe, J.P. and Haydock, K.P. (1971). Some factors affecting the growth of Lotononis (Lotononis bainesii). Australian Journal of Experimental Agriculture and Animal Husbandry, 11: 29-34.

Byth, D.E. (1964). Breeding system and chromosome number in Lotononis bainesii Baker. Nature, 202:830-831.

Cameron, D.G. (1985). Tropical and subtropical pasture legumes. 6. Lotononis (Lotononis bainesii): a very useful but enigmatic legume. Queensland Agricultural Journal 111: 2, 69-72.

Catchpoole, V.R. (1970). Laboratory ensilage of three tropical pasture legumes – Phaseolus atropurpureus, Desmodium intortum and Lotononis bainesii. Australian Journal of Experimental Agriculture and Animal Husbandry. Vol 10: 568-576.

Cook, B.G., Pengelly, B.C., Brown, S.D., Donnelly, J.L., Eagles, D.A., Franco, M.A., Hanson, J., Mullen, B.F., Partridge, I.J., Peters, M. and Schultze-Kraft, R. (2005). Tropical Forages: an interactive selection tool. [CD-ROM], CSIRO, DPI & F (Qld), CIAT and ILRI, Brisbane, Australia

Dalla Rizza, M., Real, D., Quesenberry, K.H. and Albertini, E. (2004). Plant reproductive system determination under field conditions based on co-dominant markers. Journal of Genetics and Breeding. Vol 1 47-57.

Gates, C.T. (1973). Comparative efficiency of development under cold stress of the tropical legumes Lotononis bainesii Baker and Stylosanthes humilis H.B.K. Australian Journal of Biological Science 26: 693-704.

Hutton, E. M. 1960. Flowering and Pollination in Indigofera spicata, Phaseolus lathyroides, Desmodium unicinatum, and some other tropical pasture legumes. Empire Journal of Experimental Agriculture 28 (111): 235-243.

Jaftha, J.B., Strijdom, B.W. and Steyn, P.L. (2002). Characterization of pigmented Methylobacterium bacteria which nodulate Lotononis bainesii. Systematic and Applied Microbiology 25: 440-449.

Javier, R.R. (1985). Effects of flooding on seven species of tropical pasture legumes. Annals of Tropical Research 5: 2, 91-101.

Jones, R.J., Davies, J.G. and Waite, R.B. (1967). The contribution of some tropical legumes to pasture yields of dry matter and nitrogen at Samford, southeastern Queensland. Australian Journal of Experimental Agriculture and Animal Husbandry, 7: 56-65.

Ludlow, M.M. and Wilson, G.L. (1970). Growth of some tropical grasses and legumes at two temperatures. Journal of Australian Institute of Agricultural Science, 36: 43-45.

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Note: The author acknowledges the following researchers who contributed to this profile: Mr. Mauro Zarza, Dr. Marco Dalla Rizza, Mr. Rafael Reyno, Mr. Diego F. Risso, Mr. Robin Cuadro, Dr. Nora Altier, Mr. Ignacio Risso and Mr. Andrés Martínez (Uruguay); Prof. Ken. H. Quesenberry and Dr. Mary Williams (USA); Prof. John Howieson and Dr. Ron Yates (Australia); Prof. Ben-Erik van Wyk (South Africa).It should be noted that this profile is based on the original material in Skerman et al. (1988), which has been much modified to reflect recent findings.